Dissertation

DISSERTATION

Titel der Dissertation

Comparative analysis of high-alpine and subnival vegetation of mountain ecosystems in Iran (Alborz and NW-Iran) and assessing the impacts of climate change and land-use

verfasst von Jalil Noroozi Eshlaghi

angestrebter akademischer Grad Doctor of Philosophy (PhD)

Wien, 2013

Studienkennzahl lt. Studienblatt: A 094 437 Dissertationsgebiet lt. Studienblatt: Biologie Betreut von: Univ.-Prof. Dr. Georg Grabherr

Acknowledgment I would like to express my sincere thanks to my supervisor, Prof. Georg Grabherr, who gave me the opportunity to come to Austria for my PhD study at the University of Vienna and work within the GLORIA long-term observation programme. I am deeply grateful to Harald Pauli who assisted my thesis and helped me scientifically and morally during deferent processes of the thesis. Wolfgang Willner is particularly acknowledged for supervising the vegetation task of my thesis and spending a long time to help me to publish the vegetation papers. I thank all members of GLORIA team in Vienna (Michael Gottfried, Christian Klettner, Sonya Laimer, Andrea Lamprecht and Sophie Niessner) and also members of Faculty Centre of Biodiversity for their help and reception through these years. Ernst Vitek and Walter Till, the curators of the Herbaria W and WU, respectively, are acknowledged for their help to access the plant specimens from my study area. I thank Norbert Sauberer (Gramineae), Prof F. Ehrendorfer (Rubiaceae), Prof. D. Podlech (Astragalus), Adolf Polatschek (Erysimum), Bruno Wallnöfer (Carex) and Hildemar Scholtz (Festuca and Bromus) for determination of certain critical taxa. Many thanks go to Stefan Schindler for his helps during my accommodation in Vienna. I am deeply obliged to my family and specially my wife for their supports. This work was mainly supported by the GLORIA co-ordination at the University of Vienna and the Austrian Academy of Sciences (www.gloria.ac.at).

I Table of contents

List of original publications ………………………………………………….…… 1

Author’s contribution to each publication………………………………………….... 1

Abstract………………………………………………………..…………………… 2

Zussamenfassung……………………………………...………………………..….. 4

Introduction………………………………………….……………..……………… 6

Chapter 1: Plant diversity and phytogeography at the upper limit of vascular plants of Iran and potential effect of global warming on these habitats…….…………….. 18

Chapter 2: Unknown elements of the subnival-nival flora of Iran ………………. 50

Chapter 3: Vegetation of the alpine and subnival-nival zones of N and NW Iran... 92

Conclusions……………………………………………………..………………... 137

Curriculum Vitae ……………………………………………………………….. 140

II PhD Thesis of Jalil Noroozi Eshlaghi 2013

List of original publications

The thesis is based on the following publications:

1. Noroozi, J., Pauli, H., Grabherr, G. & Breckle, S.W. (2011) The subnival-nival vascular plant species of Iran: a unique high-mountain flora and its threat from climate warming. Biodiversity and Conservation 20: 1319-1338. 2. Noroozi, J., Willner, W., Pauli, H. & Grabherr, G. (2013) Phytosociology and ecology of the high-alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.). Applied Vegetation Science DOI: 10.1111/avsc.12031. 3. Noroozi, J., Ajani, Y. (2013) A new alpine species of Nepeta sect. Capituliferae (Labiatae) from Northwestern Iran. Novon 22: 297–303. 4. Noroozi, J., Ajani, Y., Nordenstam, B. (2010) A new annual species of Senecio (Compositae- Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area. Compositae News Letter 48: 43-62. 5. Ajani, Y., Noroozi, J. & Levichev, I.G. (2010) Gagea alexii (Liliaceae), a new record from subnival zone of southern Iran with key and notes on sect. Incrustatae. Pakistan Journal of Botany 42: 67-77. 6. Noroozi, J. (In press). High mountain regions of Iran (chapter 7.6), in C. Hobohm (editor). Endemism in Vascular Plants. Springer.

Author’s contribution to each publication

Publications 1 2 3 4 5 6

Idea and design + + + + - + Sampling + + + + + + Data analysis + + + + - + Writing + + + + + +

1 PhD Thesis of Jalil Noroozi Eshlaghi 2013

Abstract

Iran is a mountainous country where rather small patches of alpine habitats are scattered across large mountain areas. The diverse flora and vegetation of these habitats have been poorly investigated so far, despite the high rate of narrowly distributed plant species which are expectedly very vulnerable to climate change impacts. The aim of this thesis is to study the flora and vegetation of the high alpine to subnival-nival elevation belts of Iran with emphasis on N- (Alborz) and NW-Iran (Sahand and Sabalan), its biodiversity in relation to elevation and its vegetation patterns in comparison with neighboring mountain systems, as well as its potential threats through climate warming. The thesis is divided into two related research foci, the first on high-altitude species, the second on their vegetation patterns. The first study deals with assessing the plant diversity and phytogeography of vascular plants occurring in the uppermost elevation belts of Iran, and attempts to give a first estimate on the risk of biodiversity losses through effects of climate change. This work was based on an extensive literature research and on additional field observations. All vascular plant species living in the subnival–nival zone of Iranian mountains (151 species) and those restricted to this zone (51) were identified. The elevational and geographical distribution patterns of these species were analyzed and the current distribution patterns are discussed with respect to potential warming- induced species losses. The rate of endemics (endemic to Iran, in most cases however far more restricted distributions) increases sharply with increasing elevation. The narrow distribution of most of Iran’s cold-adapted mountain flora and the low potential of alternative cold habitats render it highly vulnerable to climate change. Furthermore, Iran’s high mountains may host a number of still unknown species, as this study revealed two subnival-nival vascular plant species new to science: Senecio subnivalis (on Hezar Mts in south Iran) and Nepeta sahandica, (on Sahand, NW Iran), and a highly disjunct occurrence of Gagea alexii. The original descriptions of these discoverings and of these species’ vegetation ecology are arranged in a separate chapter. The second main research focus of the thesis was on the vegetation of the alpine and subnival- nival zones of N and NW Iran. In total, about 700 vegetation relevés have been collected from different habitats of the study area during the MSc and PhD theses. This thesis mainly deals with

2 PhD Thesis of Jalil Noroozi Eshlaghi 2013

high-elevation scree vegetation, using 141 relevés for analysis and phytosociological classification. A synoptic table of the most important alliances of scree vegetation in the European Alps, the Balkan Peninsula, the Caucasus, Anatolia, as well as the alliances described in the present paper was prepared to show the floristic and syntaxonomic relationship of these regions. All high-alpine and subnival scree communities of the study area were arranged in one class, two orders, and three alliances which are new for the science. This was the first formal syntaxonomic classification of the high mountain scree vegetation of Iran.

3 PhD Thesis of Jalil Noroozi Eshlaghi 2013

Zusammenfassung

Iran ist ein Gebirgsland. Alpine Habitate sind jedoch über ausgedehnte Gebirgsräume weit zerstreut und stark fragmentiert. Die artenreiche Flora und Vegetation dieser Habitate wurde bislang kaum untersucht. Dies trotz des hohen Anteils engräumig verbreiteter Arten und der dadurch erwarteten hohen Anfälligkeit auf Effekte des Klimawandels. Ziel dieser Dissertation ist das Studium der Flora und Vegetation der hoch-alpinen bis subnival- nivalen Höhenstufen Irans, mit einem speziellen Fokus auf das Elburs-Gebirge in Nord-Iran und die isolierteren Bergstöcke Sahand und Sabalan im Nordwesten. Im speziellen bezieht sich die Arbeit auf Biodiversitätsmuster entlang des Höhengradienten, auf die Vegetationszusammensetzung im Vergleich mit benachbarten Gebirgsräumen und auf das Gefährdungspotential bezüglich Klimaerwärmung. Die Arbeit ist in zwei Themenbereiche gegliedert, wobei sich der erste auf die Artenmuster, der zweite auf die Vergesellschaftung der Arten bezieht. Teil 1 beinhaltet eine detaillierte phytogeographische Erfassung aller in den obersten Höhenstufen vorkommenden Gefäßpflanzenarten und versucht, darauf aufbauend, eine erste Risikoabschätzung von klimawandelinduzierten Biodiversitätsverlusten zu geben. Die Studie basiert auf eingehender Literaturrecherche sowie ergänzenden Felderhebungen. Für die gesamte subnival-nival-Stufe wurden 151 Arten identifiziert, 51 davon sind auf diese Höhenstufe beschränkt. Anhand der Verbreitungsmuster der Arten entlang des Vertikalgradienten und der geographischen Verbreitung wird das Potential für erwärmungsinduzierte Artenverluste diskutiert. Der Endemitenanteil (Iran-Endemiten, meist jedoch sind es sehr lokal verbreitete Arten) zeigte einen sehr deutlichen Anstieg mit der Höhenlage. Sehr engräumig verbreitete Arten zeigen also eine Häufung in der höchsten, nur sehr kleinflächigen Höhenstufe. Diese kälteadaptierten Gebirgsspezialisten sind demnach hochgradig anfällig auf Klimawandeleinflüsse. Darüber hinaus sind für die iranischen Gebirge noch etliche unbekannte Gefäßpflanzenarten zu erwarten, wie durch die Neufunde im Rahmen dieser Studie angezeigt ist. Zwei Arten, Senecio subnivalis (im Hezar-Gebirge Süd-Irans) und Nepeta sahandica (im Sahand, NW-Iran), wurden erstmals beschrieben, für eine weitere, Gagea alexia, wurde ein sehr disjunktes Vorkommen gefunden. Die Originalbeschreibungen mit vegetationsökologischen Informationen zu diesen Arten sind in einem eigenen Kapitel dargestellt.

4 PhD Thesis of Jalil Noroozi Eshlaghi 2013

Der zweite Haupt-Fokus der Dissertation liegt auf der Vegetation der hoch-alpinen und subnival- nivalen Stufe Nord- und Nordwest-Irans. Aus insgesamt 700 Vegetationsaufnahmen wurden 141 Aufnahmen für die Analyse und phytosoziologische Klassifizierung der Schuttvergesellschaftungen ausgewählt. Anhand einer synoptischen Tabelle wurde der syntaxonomische Bezug mit der Schuttvegetation benachbarter Gebirge in Anatolien, Kaukasus sowie der Gebirge auf der Balkanhalbinsel und der Alpen untersucht. Aufgrund klarer floristischer Differenzierung zu den Nachbargebirgen mussten die iranischen Schuttgesellschaften in eine neue syntaxonomische Klasse mit zwei Ordnungen und drei Verbänden gestellt werden. Die Arbeit stellt die erste formale syntaxonomische Klassifikation der Hochgebirgs-Schuttgesellschaften Irans dar.

5 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

Introduction

1 Background

Mountains are hot spots of biodiversity related to the steep elevational gradients and, thus, to a compression of bioclimatic zones, to topographic heterogeneity, and to speciation through isolation (Nagy & Grabherr 2009). The alpine life zone in the plant geographic sense is applied to any low stature vegetation above the climatic treeline worldwide, and alpine vegetation represents the only biogeographic unit on land with a global distribution and is fragmented into many mountain regions (Körner 2003). The alpine floras of the world are nested within a great variety of regional floras, partly explaining the great overall species diversity among alpine vegetation (Körner & Spehn 2002). Different phenomena in high mountains such as geographic isolation, tectonic uplift, climatic changes, glaciation, strong micro habitat differentiation and a varied history of migration and/or evolution cause a high rate of taxonomic richness (Packer 1974; Agakhanyanz & Breckle 1995; Körner 1995). Many alpine regions are biodiversity hotspots and habitat for many endemic species (Körner 2003). Alpine vegetation and the distribution ranges of plant species are affected by climate change. Studies, however, are mainly available for temperate and boreal mountains (Grabherr et al. 1994; Walther et al. 2005a; Pauli et al. 2007; Vittoz et al. 2008; Klanderud & Birks 2003; Britton et al. 2009). Resurveys in alpine areas of a number of European mountain systems revealed that vascular plants have been migrating upward to higher altitudes (Grabherr et al. 1994, 2001; Pauli et al. 1996, 2007, 2012; Sturm et al. 2001; Moiseev & Shiyatov 2003; Walther et al. 2005a, 2005b; Holzinger et al. 2008; Klanderud & Totland 2005; Parolo & Rossi 2008; Erschbamer et al. 2009; Gottfried et al. 2012). Biota of high mountains around the Mediterranean Basin including Iran must be considered as being exceptionally vulnerable regarding climate warming impacts due to the small and isolated alpine and subnival-nival areas and due to the potential of increasing drought stress through climate change (Pauli et al. 2003). These mountains are characterised by a pronounced seasonality of precipitation patterns, with dry summers and precipitation in winter that falls almost exclusively as snow at high elevations. A precondition for the exploration and the assessment of potential affects of climate change but also for the interpretation of future monitoring data, is a thorough assessment of the flora and vegetation. For the Iranian alpine life zone, climate impact studies are completely lacking and vegetation studies

6 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

are very scarce. Therefore, alpine and subnival vegetation ecology and phytosociological description was one of the core components of my thesis. Vegetation ecology, the study of the plant cover and its relationships with the environment, is an intricate scientific work, both due to the overwhelming variation of its object of study in space and time, and its complicated interactions with abiotic and biotic factors (van der Maarel 2005). Phytosociology is a part of vegetation science, with focusing on extant, taxonomic plant assemblages at the scale of vegetation stands, and its essential purpose is the definition and functional characterization of vegetation types based on the total floristic composition of stands (Dengler et al. 2008). A consistent large-scale classification of floristically defined vegetation types is an important tool for ecological research, bioindication, vegetation monitoring, conservation strategies and legislation (Dengler et al. 2008).

2 Alpine habitats of Iran

Iran with a total surface area of c. 1.6 km², is mostly a high plateau in the SW Asia. Almost half of the country is structured with high elevations and interior lowlands are surrounded with high mountains including the Alborz Mountains in the north, Zagros Mountains extending from the the northwest to southeast, Azerbaijan Plateau in the northwest, Kopeh-Dagh Mountains in the northeast and Makran Mountains in southeast (Fig. 1). Alpine habitats are almost above 3000 m a.s.l. and scattered in different parts of country. In these ecosystems, the summer is arid, hot and sunny, with intense radiation most of the time. Both annual and diurnal amplitudes of temperature can be very high, in particular near the soil surface. The high elevations of Alborz are affected by the north-westerly flow of polar air masses, and annual precipitation reaches almost 1000 mm (Khalili 1973). In total c. 700 vascular plant species are present in the alpine belt of Iranian mountains, where more than 50% are endemic or subendemic to the country (Noroozi et al. 2008). The alpine flora of Iran is mainly of Irano-Turanian origin (Zohary 1973; Klein 1982, 1991; Frey et al. 1999; Noroozi et al. 2008). It exhibits the strongest floristic relationships with Anatolia, Hindu Kush and Caucasus which have 23%, 20% and 19% species in common with the high mountains of Iran, respectively (Noroozi et al. 2008).

7 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

The N- and NW-Iran is known as being particularly rich in species including many endemics, but ecological studies on the region’s alpine plants are scarce or almost absent in the scientific literature. Phytosociological studies of the alpine zone of Iran were carried out only in Central Alborz, and the syntaxonomic inventory of Iran is still far from being complete. The studies of Bunge (1960), Kotschy (1861a, b), Buhse (1899a, b) and Gilli (1939, 1941) were the initial vegetation surveys on high altitudes of Central Alborz. Klein (1991) published a paper on endemism of alpine flora of Alborz mountains. An overview of the flora and phytogeography of the alpine zone in the Iranian mountains was provided by Noroozi et al. (2008). The most important phytosociological research in the subalpine, alpine and subnival belt which aimed at establishing a syntaxonomic system was carried out by Klein (1982, 1984, 1987, 1988, 1994) and Klein & Lacoste (1994, 1996, 1998, 2001). Noroozi et al. (2010) presented a new classification of the snowbed and thorn-cushion grasslands of the alpine zone of Central Alborz. A synopsis of plant communities and their elevational patterns in the southern slopes of Central Aborz from the low montane to the high alpine zone are presented in Akhani et al. (2013).

Fig. 1. Mountain ranges of Iran and their altitudinal zonation.

8 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

3 Aims of the study

This dissertation thesis focused on the high mountain flora and vegetation of Iran with emphasize on N- (Alborz) and NW-Iran (Sabalan and Sahand), its biodiversity in relation to altitude and to the neighboring mountain systems, and its potential threats through climate warming.

3.1 The research questions 1) How many vascular plant species live in Iran’s subnival–nival zone? 2) Are there species being restricted to the subnival–nival zone? 3) What is the relationship between their altitudinal and geographical distribution patterns and are the latter different from species dwelling at lower elevations? 4) Which threats and implications for conservation biology may arise for narrowly distributed cryophilic species? 5) Which major plant community types can be distinguished in high-altitude screes in Central Alborz and the mountains of NW Iran on the basis of their compositional variation, as well as on species’ ecological, life-form, and biogeographical characteristics and their habitat preferences? 6) What are the syntaxonomic relationships between these vegetation types at similar habitats of

Anatolia, the Caucasus, the Balkan Peninsula and the European Alps?

3.2 Hypotheses 1) Mediterranean high mountains of southern Europe are highly isolated from each other, remained partly ice-free during the Pleistocene and, therefore, show a high rate of endemism, particularly at high elevations (Pauli et al. 2003; Stanisci et al. 2005). Given that the high mountains of Iran share similar geomorphological, glaciation and climatic features, we hypothesize that the majority the vascular plant species in Iranian high mountains are endemics (species restricted to particular mountain ranges or at least to the country) and that their proportion is increasing with elevation. 2) Alpine regions in Iran are scattered over a large area and floristic as well as vegetation studies are largely lacking or only were conducted very sporadically. We, therefore, expect to find new phytogeographic details of known species and probably not yet described species. 3) If the majority of high-altitude species, in fact, are endemics, we hypothesize that plant communities in Iranian mountains belong to different phytosociological units as previously

9 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

described communities in similar habitats of the adjacent regions (Anatolia, Caucasus, Balkan and Alps).

4 Results

The result of this thesis is organized into three major chapters: 1) Plant diversity and phytogeography at the upper limit of vascular plants of Iran and potential effect of global warming on these habitats 2) Unknown elements of the subnival-nival flora of Iran 3) Vegetation of the alpine and subnival-nival zones of N and NW Iran

4.1 Chapter 1: Plant diversity and phytogeography at the upper limit of vascular plants of Iran and potential effect of global warming on these habitats A paper and a book chapter were published for this chapter:

4.1.1 Paper: Noroozi, J., Pauli, H., Grabherr, G. & Breckle, S.W. (2011) The subnival-nival vascular plant species of Iran: a unique high-mountain flora and its threat from climate warming. Biodiversity and Conservation 20: 1319-1338. This paper provides a first country-wide overview of the vertical distribution patterns and the chorology of vascular plant species that occur in the subnival-nival zone of Iran. After the paper of Noroozi et al. (2008) on alpine habitats of Iran, which represents plant diversity and phyogeography of these ecosystems, it was worthful to focus in detail on phytodiversity of the upper limit of vascular plants which are more cold-adapted and probably more vulnerable to ongoing global warming. In this work we showed how many plant species are living in subnival- nival zone and how many of them are restricted to these habitats. The elevational and geographical distribution patterns of these species were classified and discussed. A total of 151 vascular plant species were identified to occur in the subnival–nival zone of Iran, where 51 of these can be considered as true subnival–nival species. Within the latter, the degree of species endemic to Iran is 68% and clearly decreases to 53 and 20% for species that also occur in the alpine and in the subalpine zones, respectively. The rate of endemism increases very sharply with increasing altitude in Iran. Furthermore, a narrow vertical distribution restricted to high altitudes

10 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

often corresponds to a constricted geographic range. Vice versa, species with a larger amplitude of their elevational distribution commonly are widespread species. Richness and proportions of endemic species, in fact, were found to be similar to mountains around the Mediterranean Basin, such as the central Apennines (Stanisci et al. 2005), Sierra Nevada (Pauli et al. 2003) or the Atlas mountains (Favarger 1972). The larger alpine areas further north in Europe, such as the Alps, host fewer endemic species, which predominantly occur in the out and lower ranges, where glaciation in the ice ages was weak (Merxmüller 1952; Essl et al. 2011). In the Caucasus mountains the percentage of endemics was reported to be high in the subnival belt (Nakhutsrishvili 1998), but further comparative studies along elevation gradients would be desirable. In a larger, northern hemisphere view, the outstandingly high degree of high-elevation endemism is not a general feature, but appears to be especially pronounced in the mountain ranges exposed to Mediterranean-type climate west of the Hindu Kush to NW-Africa and SW-Europe. In the large and more connected mountain ranges of the Hindu Kush-Himalaya-Central Asia system endemism is less common. In the Hindu Kush, for example, endemism peaks in the subalpine zone, but species dwelling in the nival zone (above 5000 m) mostly have a wide geographical distribution over the Eurasian mountains and some to the Arctic (Breckle 1974, 2004). Also in most of the large mountain systems of North America, the degree of endemism is rather low (Mills & Schwartz 2005), which seems to be related to the availability of effective migration pathways in these north-south-oriented mountain ranges. The exceptionally high level of high- altitude endemism appears to result mainly from a pronounced current orographic isolation and fragmented cold areas and from the absence of extensive Pleistocene glaciations. The narrow distribution of most of Iran’s cold-adapted mountain flora and the low potential of alternative cold habitats render it highly vulnerable to climate change.

4.1.2 Book chapter: Noroozi, J. (in press) High mountain regions of Iran (chapter 7.6), in C. Hobohm (editor). Endemism in Vascular Plants. Springer. In this accepted manuscript the rare and narrow distributed alpine species of Iranian mountains which are known only from one or very few locations are introduced. A total of 110 vascular plant species were considered to be rare and narrow endemic to Iranian alpine habitats. These species were found mainly in Zagros, Alborz, and in the northwestern part of the country. The species were classified according to their geographical distributions. Most of them can be

11 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

categorized as Endangered (EN) and Critically Endangered (CR) according to IUCN Red List criteria. In this work the relationship of rate of endemism and different habitats in alpine zone is discussed.

4.2. Chapter 2: Unknown elements of the subnival-nival flora of Iran Due to the low state of research, high-altitude botanical investigations still yield new species for Iran and to science. The following three papers are original descriptions of two new species, and of one species new to Iran, all occurring in the subnival-nival elevation belt of Iran:

4.2.1 Paper 1: Noroozi, J., Ajani, Y., Nordenstam, B. (2010) A new annual species of Senecio (Compositae- Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area. Compositae News Letter 48: 43-62. In this paper, Senecio subnivalis was described as a new species for science from the subnival zone of Hezar mountain (southern Iran). The accompanying species of the new species and some ecological features of its habitat were presented. Moreover, the phytogeographical aspects of the Hezar-Lalehzar Mts. in high elevations were discussed and different distribution patterns of the species were presented.

4.2.2 Paper 2: Ajani, Y., Noroozi, J. & Levichev, I.G. (2010) Gagea alexii (Liliaceae), a new record from subnival zone of southern Iran with key and notes on sect. Incrustatae. Pakistan Journal of Botany 42: 67-77. In this paper, Gagea alexii was recorded for the first time for Iran from subnival zone of Hezar Mt., exactly in the habitat of Senecio subnivalis. It has a very long disjunct distribution from Central Asia and northern Hindu Kush mountains, with a large gap to the isolated occurrence in southern Iran. This seems to be the result of post-glacial global warming. The occurrence of this species in the subnival zone of the high mountains of south Iran is a further evidence of the close floristic affinity of the southeastern Zagros with the Hindu Kush and Central Asia, especially at high altitudes. In this paper the taxonomic problems of the genus Gagea in Iran is also discussed.

4.2.3 Paper 3: Noroozi, J., Ajani, Y. (2013) A new alpine species of Nepeta sect. Capituliferae (Labiatae) from Northwestern Iran. Novon 22: 297–303.

12 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

In this paper Nepeta sahandica was described and illustrated from the high elevations of the Sahand Mountains in northwestern Iran. Morphological differences of this species from its closest relatives were discussed. The new species is restricted to the subnival vegetation zone, associated with unstable scree grounds. The associated species of Nepeta sahandica and ecological aspects of its habitat were presented in a phytosociological table. This species was classified as Critically Endangered (CR), based on IUCN Red List criteria.

4.3 Chapter 3: Vegetation of the alpine and subnival-nival zones of N and NW Iran One paper, representing one of the core publications of the thesis, was published for this chapter:

4.3.1 Paper: Noroozi, J., Willner, W., Pauli, H. & Grabherr, G. (2013) Phytosociology and ecology of the high alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.). Applied Vegetation Science DOI: 10.1111/avsc.12031. As discussed in Noroozi et al. (2010), the phytosociological classification of Klein (1982, 1987, 1988) and Klein & Lacoste (2001) from alpine and subnival zones of Iran are not convincing and need to be revised. The paper of Noroozi et al. (2010) mainly focused on the ecology and phytosociology of snowbed and thorn-cushion grassland vegetation types of alpine zone of Central Alborz and described and classified different plant communities. It was a later addition to the master thesis of the author, but major parts of analyses and the writing process were carried out during the PhD thesis in Vienna. After this paper, the priority focus for vegetation analysis was on scree vegetation types of alpine and subnival-nival zones of the study area. A total of 141 phytosociological relevés from 3200 up to 4800 m a.s.l. of Alborz and Azerbaijan mountains have been collected and used in this paper including 14 relevés of Klein & Lacoste (2001) from similar habitats of Central Alborz. This data set was classified using the TWINSPAN algorithm, and the numerical classification was translated into a syntaxonomic system. In this work, we described one class (Didymophyso aucheri-Dracocephaletea aucheri), two orders (Physoptychio gnaphalodis-Brometalia tomentosi, Didymophysetalia aucheri), three alliances (Elymo longearistati-Astragalion macrosemii, Erigerontion venusti, Didymophysion aucheri) and 10 associations. The ecological characters of the all communities were presented and the geographical territory of them discussed. The territory of this class, which is the first valid phytosociological class of the high elevations of Iran, extends from Alborz to NW Iran and

13 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Introduction

probably to E Anatolia, Transcaucasia and to the Zagros mountains. A synoptic table of the most important alliances of scree vegetation in the European Alps, the Balkan Peninsula, the Caucasus, Anatolia, as well as the alliances described in the present paper was prepared to show the floristic and syntaxonomic relationship of these regions.

References

Agakhanjanz, O. & Breckle S-W. (1995) Origin and evolution of the mountain flora in middle Asia and neighboring mountain regions. In: Chapin III FS, Körner C (eds) Arctic and alpine biodiversity: patterns, causes and ecosystem consequences, pp. 63–80. Ecological Studies 113, Springer, Berlin, DE. Akhani, H., Mahdavi, P., Noroozi, J. & Zarrinpour, V. (2013) Vegetation pattern of Irano- Turanian steppes along a 3000 m altitudinal gradient in the Alborz Moutains (Iran). Folia Geobotanica 48: 229-255. Breckle, S-W. (1974) Notes on Alpine and Nival Flora of the Hindu Kush, East Afghanistan. Botaniska Notiser (Lund) 127:278–284. Breckle, S-W. (2004) Flora, Vegetation und Ökologie der alpin–nivalen Stufe des Hindukusch (Afghanistan). In: Breckle, S-W., Schweizer, B. & Fangmeier, A. (eds) Proceedings of 2nd symposium AFW Schimperfoundation: results of worldwide ecological studies. Stuttgart- Hohenheim, p 97. Britton, A.J., Beale, C.M., Towers W. & Hewison R.L. (2009) Biodiversity gains and losses: Evidence for homogenisation of Scottish alpine vegetation. Biological Conservation 142: 1728-1739. Buhse, F. (1899a) Flora des Alburs und der Kaspischen Südküste. Bisherige Forschungsergebnisse aus diesem Gebiet. Arbeiten des Naturforscher-Vereins zu Riga. Neue Folge. 8. Heft. Buhse, F. (1899b) Reisebemerkungen aus dem östlichen Albursgebirge in Persien. Bulletin de la Société Impériale des Naturalistes de Moscou 34: 363-383. Bunge, A. (1860) Die Russische Expedition nach Chorassan in den Jahren 1858 und 1859. Petermanns Geographische Mitteilungen 6: 205–226. Dengler, J., Chytrý, M. & Ewald, J. (2008) Phytosociology. In: Jørgensen, S. E. & Fath, B. D. (eds.) Encyclopedia of ecology, pp. 2767–2779. Elsevier, Oxford, UK. Erschbamer, B., Kiebacher, T., Mallaun, M. & Unterluggauer, P. (2009) Short-term signals of climate change along an altitudinal gradient in the South Alps. Plant Ecology 202: 79-89. Essl, F., Dullinger, S., Plutzar, C, Willner, W. & Rabitsch, W. (2011) Imprints of glacial history and current environment on correlations between endemic plant and invertebrate species richness. Journal of Biogeography 38: 604–614.

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Favarger, C. (1972) Endemism in the montane floras of Europe. In: Valentine DH (ed) Taxonomy, phytogeography and evolution. Academic Press, London, pp 191–204. Frey,W., Kürschner, H. & Probst, W. (1999) Flora and vegetation, including plant species and larger vegetation complexes in Persia. In: Yarshater, E. (ed.) Encyclopaedia Iranica 10(1), pp. 43–63. Mazda Publishers, Costa Mesa, CA, US. Gilli, A. (1939) Die Pflanzengesellschaften der Hochregion des Elbursgebirges in Nordiran. Beiheft des Botanischen Centralblatts 59: 317-344. Gilli, A. (1941) Ein Beitrag zur Flora des Elburs-Gebirges in Nord-Iran. Feddes Repertorium Specierum Novarum Regni Vegetabilis 50: 263-283. Gottfried, M., Pauli, H., Futschik, A., Akhalkatsi, M., Barancok, P., Benito Alonso, J. L., Coldea, G., Dick, J., Erschbamer, B., Fernández Calzado, M. R., Kazakis, G., Krajci,J., Larsson, P., Mallaun, M., Michelsen, O., Moiseev, D., Moiseev, P., Molau, U., Merzouki, A., Nagy, L., Nakhutsrishvili, G., Pedersen, B., Pelino, G., Puscas, M., Rossi, G., Stanisci, A., Theurillat, J.- P., Thomaselli, M., Villar, L., Vittoz, P., Vogiatzakis, I. & Grabherr, G. (2012) Continent- wide response of mountain vegetation to climate change. Nature climate change 2: 111-115. Grabherr, G., Gottfried, M. & Pauli, H., (1994) climate effects on mountain plants. Nature 369: 448-448. Grabherr, G., Gottfried, M. & Pauli, H. (2001) Long-term monitoring of mountain peaks in the Alps. In: Burga, C.A., & Kratochwil, A. (eds.) Biomonitoring: General and Applied Aspects on Regional and Global Scales, Vol 35, pp 153-177. Tasks for Vegetation Science, Kluwer, Dordrecht. Holzinger, B., Hülber, K., Camenisch, M. & Grabherr, G. (2008) Changes in plant species richness over the last century in the eastern Swiss Alps: elevational gradient, bedrock effects and migration rates. Plant Ecology 195:179–196. Khalili, A. (1973) Precipitation patterns of Central Elburz. Archiv für Meteorologie, Geophysik und Bioklimatologie, Serie B 21: 215–232. Klanderud, K. & Birks, H.J.B. (2003) Recent increases in species richness and shifts in altitudinal distributions of Norwegian mountain plants. The Holocene 13: 1-6. Klanderud, K, Totland, Ø (2005) Simulated climate change altered dominance hierarchies and diversity of an alpine biodiversity hotspot. Ecology 86: 2047–2054. Klein, J.C. (1982) Les groupements chionophiles de l’Alborz central (Iran). Comparaison avec leurs homologues d’ Asie centrale. Phytocoenologia 10: 463–486. – (1984) Les groupements végétaux d’altitude de l’Alborz central (Iran). – Ecologie des milieux montagnards et de haute altitude. Documents d’Ecologie Pyrénéenne 3–5: 199–204. – (1987) Les pelouses xérophiles d’altitude du flanc sud de l’Alborz central (Iran). Phytocoenologia 15: 253–280. – (1988) Les groupements à grandes ombellifères et à xérophytes orophiles: Essai de synthèse à l’échelle de la région irano-touranienne. Phytocoenologia 16: 1–36. – (1991) Endémisme à l’étage alpin de l’Alborz (Iran). Flora et Vegetatio Mundi 9: 247–261.

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– (1994) La végétation altitudinale de L’Alborz central (Iran). Institut Français de Recherche en Iran (Téhéran). Klein, J.C. & Lacoste, A. (1994) Les pelouses subalpines (Alchemilletum plicatissimae ass. nov.) de l’Alborz central (Iran): ultime avancée sud-orientale de l’aire des Festuco-Brometea Br.-Bl. et Tx. 1943. Phytocoenologia 24: 401–421. – (1996) Aperçu synthétique sur l’ étagement de végétation dans l’Alborz central (Iran). Annalen des Naturhistorischen Museums in Wien 98: 67–74. – (1998) L‘étagement de végétation dans le massif de l‘Alborz central (Iran): essai de comparison avec celui du systéme alpin et des montagnes méditerranéennes. Ecologie 29: 181–186. – (2001) Observation sur la végétation des éboulis dans les massifs irano-touraniens: le Galietum aucheri ass. nov. de l’ Alborz central (N-Iran). Documents Phytosociologiques N. S. 19: 219- 228. Kotschy, T. (1861a) Die Vegetation des westlichen Elbrus in Nordpersien. Österreichische Botanische Zeitschrift 11: 105-117. Kotschy, T. (1861b) Der westliche Elbrus bei Teheran. Mitteilungen der Kaiserlich-Königlichen Oestereichischen Geographischen Gesellschaft Wien 5: 65–110. Körner, Ch. (1995) Alpine plant diversity: a global survey and functional interpretations. In: Chapin FS III, Körner Ch (eds.) Arctic and alpine biodiversity: Patterns, causes and ecosystem consequences, pp. 45-62. Springer Ecol Studies 113, Berlin, Heidelberg, New York. Körner, Ch. (2003) Alpine Plant Life. Functional Plant Ecology of High Mountain Ecosystem. Springer, Berlin. Körner, Ch., & Spehn E.M. (eds.) (2002) Mountain biodiversity. A global assessment. Parthenon, Boca Raton. Merxmüller, H. (1952) Untersuchungen zur Sippengliederung und Arealbildung in den Alpen, Teil I. Jahrb Ver Schutze Alpenpflanzen Tiere 17:96–133. Mills, M-H., Schwartz, M-W. (2005) Rare plants at the extremes of distribution: broadly and narrowly distributed rare species. Biodiversity and Conservation 14: 1401–1420. Moiseev, P.A. & Shiyatov S.G. (2003) Vegetation dynamics at the treeline ecotone in the Ural highlands, Russia. In: Nagy, L., Grabherr, G., Körner, C. & Thompson D.B.A. (eds.) Alpine biodiversity in Europe – A Europe-wide assessment of biological richness and change, pp 423-435. Springer, Berlin, DE. Nakhutsrishvili, G. (1998) The vegetation of the subnival belt of the Caucasus Mountains. Arctic and and Antarctic Alpine Research 30: 222–226. Noroozi, J., Akhani, H. & Breckle, S.W. (2008) Biodiversity and phytogeography of the alpine flora of Iran. Biodiversity and Conservation 17: 493-521. Noroozi, J., Akhani, H. & Willner, W. (2010) Phytosociological and ecological study of the high alpine vegetation of Tuchal Mountains (Central Alborz, Iran). Phytocoenologia 40: 293-321.

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Nagy, L, Grabherr, G. (2009) The biology of alpine habitats. Oxford University Press, New York. Packer, J.G. (1974) Differentiation and dispersal in alpine floras. Arctic and and Antarctic Alpine Research 6: 117-128. Parolo, G. & Rossi, G. (2008) Upward migration of vascular plants following a climate warming trend in the Alps. Basic Applied Ecology 9: 100–107. Pauli, H., Gottfried, M. & Grabherr, G. (1996) Effects of climate change on mountain ecosystems - upward shifting of alpine plants. World Resource Review 8: 382-390. Pauli, H., Gottfried, M., Dirnböck, T., Dullinger, S. & Grabherr, G. (2003). Assessing the long- term dynamics of endemic plants at summit habitats. In: Nagy, L., Grabherr, G., Körner, C. & Thompson D.B.A. (eds.) Alpine biodiversity in Europe – A Europe-wide assessment of biological richness and change, pp. 195-207. Springer, Berlin, DE. Pauli, H., Gottfried, M., Reiter, K., Klettner, C. & Grabherr, G. (2007) Signals of range expansions and contractions of vascular plants in the high Alps: observations (1994-2004) at the GLORIA* master site Schrankogel, Tyrol, Austria. Global Change Biology 13: 147-156. Pauli, H., Gottfried, M., Dullinger, S., Abdaladze, O., Akhalkatsi, M., Benito Alonso, J.L., Coldea, G., Dick, J., Erschbamer, B., Fernández Calzado, R., Ghosn, D., Holten, J.I., Kanka, R., Kazakis, G., Kollár, J., Larsson, P., Moiseev, P., Moiseev, D., Molau, U., Molero Mesa, J., Nagy, L., Pelino, G., Puşcaş, M., Rossi, G., Stanisci, A., Syverhuset, A.O., Theurillat, J.-P., Tomaselli, M., Unterluggauer, P., Villar, L., Vittoz, P. & Grabherr, G. (2012) Recent plant diversity changes on Europe’s mountain summits. Science 336: 353-355. Stanisci, A., Pelino, G. & Blasi, C. (2005) Vascular plant diversity and climate change in the alpine belt of the central Apennines (Italy). Biodiversity and Conservation 14: 1301–1318. Sturm, M., Racine, C. & Tape, K. (2001) Climate change – increasing shrub abundance in the Arctic. Nature 411: 546-547. van der Maarel, E. (ed.) (2005) Vegetation ecology. Oxford, Blackwell, UK. Vittoz, P., Bodin, J., Ungricht, S., Burga, C. & Walther, G.R. (2008) One century of vegetation change on Isla Persa, a nunatak in the Bernina massif in the Swiss Alps. Journal Vegetation Science 19: 671-680. Walther, G-R., Beißner, S. & Burga, C.A. (2005a) Trends in the upward shift of alpine plants. Journal Vegetation Science 16: 541–548. Walther, G-R., Berger, S. & Sykes, M.T. (2005b) An ecological ‘footprint’ of climate change. Proceedings of the Royal Society of London, B 272: 1427–1432. Zohary, M. (1973) Geobotanical foundations of the Middle East, vol 2. Fischer, Stuttgart.

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Chapter 1:

Plant diversity and phytogeography at the upper limit of vascular plants of Iran and potential effect of global warming on these habitats

A paper and a book chapter were published regarding to this subject:

Paper: The subnival-nival vascular plant species of Iran: a unique high-mountain flora and its threat from climate warming

Book chapter: High mountain regions of Iran (chapter 7.6), in C. Hobohm (editor). Endemism in Vascular plants. Springer.

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Title of paper: The subnival-nival vascular plant species of Iran: a unique high- mountain flora and its threat from climate warming

Authors: Jalil Noroozi, Harald Pauli, Georg Grabherr, Siegmar-W. Breckle

Status: published (2011), Biodiversity and Conservation 20:1319–1338

Contribution: idea, data collection, data analysis, manuscript writing.

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Biodivers Conserv (2011) 20:1319–1338 DOI 10.1007/s10531-011-0029-9

ORIGINAL PAPER

The subnival–nival vascular plant species of Iran: a unique high-mountain ﬂora and its threat from climate warming

Jalil Noroozi • Harald Pauli • Georg Grabherr • Siegmar-W. Breckle

Received: 4 August 2010 / Accepted: 26 February 2011 / Published online: 10 March 2011 Ó Springer Science+Business Media B.V. 2011

Abstract This study provides a first country-wide overview of the vertical distribution patterns and the chorology of vascular plant species that occur in the uppermost elevation zones in Iran. The current distribution patterns are discussed with respect to potential warming-induced species losses. Iran’s subnival and nival vegetation zones are found at elevations above 3600–3900 m in a highly fragmented distribution across Alborz, Zagros, and NW-Iran. Based on literature research and on field observations, all vascular plant species living in the subnival–nival zone of Iranian mountains were identified (151 species) and classified into three altitudinal groups: Group A comprises species that occur mainly in subnival–nival habitats (51 species). Group B are species being common in subnival–nival areas but are equally present in the alpine zone (56 species). Group C are species that can reach to subnival areas but also grow in alpine, subalpine and sometimes lower altitudes (44 species). The chorological patterns differ among the three groups. The percentage of species being endemic to Iran decreases from group A, to B and C, with 68, 53 and 20%, respectively. A narrow altitudinal distribution at high elevations is clearly related to a small-scaled geographical distribution range. The outstanding rate of high-altitude endemism appears to result mainly from orographic isolation of the country’s highly scattered cold areas and by the absence of extensive Pleistocene glaciations. The narrow distribution of most of Iran’s cold-adapted mountain flora and the low potential of alternative cold habitats render it highly vulnerable to climate change.

J. Noroozi (&) Á G. Grabherr Department of Conservation Biology, Vegetation and Landscape Ecology, University of Vienna, Rennweg 14, 1030 Vienna, Austria e-mail: [email protected]

H. Pauli Institute of Mountain Research (IGF) of the Austrian Academy of Sciences, C/O University of Vienna, Rennweg 14, 1030 Vienna, Austria

S.-W. Breckle Department of Ecology, University of Bielefeld, Wasserfuhr 24-26, 33619 Bielefeld, Germany 123

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1320 Biodivers Conserv (2011) 20:1319–1338

Keywords Altitudinal distribution Á Climate change Á Endemism Á High mountains Á Phytogeography Á Subnival–nival zone Á Vascular plants

Introduction

Only the highest mountain peaks of Iran exceed the elevational limits of vascular plant life: Damavand (5671 m), Alamkuh (4850 m), both in the Alborz mountains and Sabalan (4811 m) in the NW of the country. The uppermost zones where vascular plants occur are the subnival and nival vegetation zones. The subnival zone is the transition zone between the alpine grassland and tragacanth heath and the scanty, patchy vegetation of the nival zone (also termed as the alpine–nival ecotone; Nagy and Grabherr 2009). The subnival– nival zone above the alpine zone has a highly scattered distribution over the major mountain systems of Iran. Its ﬂora and vegetation is less known in comparison to that of lower altitudes. Initial vegetation survey on Central Alborz that at least marginally included subnival areas dates back to Kotschy (1861a, b), Buhse (1899a, b) and Gilli (1939, 1941). Further studies on the subnival–nival vegetation of the Central Alborz were published by Klein (1982) and Klein and Lacoste (1999), who made 31 phytosociological releve´s in subnival to nival areas above 3900 m up to 4350 m. Due to the low state of research, high-altitude botanical investigation still yield new species for Iran and to science. For example, Senecio subnivalis was described most recently from Hezar–Lalehzar Mts. at 4443 m a.s.l. (Noroozi et al. 2010a). In the same place Gagea alexii was ﬁrst recorded for Iran (Ajani et al. 2010). In this study we attempt to answer the following questions: (1) how many vascular plant species live in Iran’s subnival–nival zone?; (2) what is the relationship between their altitudinal and geographical distribution patterns?; (3) are there species being restricted to the subnival–nival zone? (4) Which threats and implications for conservation biology may arise for narrowly distributed cryophilic species?

Methods

Study area

Iran is a mountainous country in Southwest-Asia covering an area of 1.6 million km2. The Alborz in the north and Zagros from northwest to southeast are the major mountain ranges of Iran. The elevation extends from -26 m a.s.l. on the shore of the Caspian Sea to 5671 m a.s.l. at the highest peak of Alborz (Damavand). The high mountains of Iran exhibit a continental climate with Mediterranean precipitation regime. The growing season is arid and exposed to intensive radiation and strong winds are common. Both annual and diurnal amplitudes of temperature can be high, on the soil surface in particular (Noroozi et al. 2008, 2010b). There are some active glaciers in both Alborz and Zagros mountain ranges (Ferrigno 1991). Subnival and nival zones have a fragmental and highly scattered distribution over the mountain systems of Alborz, NW-Iran and Zagros. The largest subnival–nival areas are concentrated in Central Alborz with more than 50 mountain peaks exceeding 4000 m. Sabalan and Sahand mountains in the northwest of the country, Zardkuh and Oshtorankuh in Central Zagros and Dena and Hezar–Lalehzar mountains in south and southeastern Zagros are other important mountains which embrace isolated subnival to nival areas (Fig. 1). 123

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Fig. 1 The mountain areas of Iran (grey) and the distribution of the subnival-nival zone (black patches)

Vertical species ranges

The documentation of species that reach subnival–nival areas and the characterization of their altitudinal ranges are based on literature, Flora Iranica in particular (Rechinger 1963– 2010), and our own ﬁeld survey. Previous phytosociological survey from Central Alborz (Klein 1982; Klein and Lacoste 1999) includes data on the region’s subnival–nival ﬂora. Our botanical surveys were carried out at different mountains of Central Alborz (Tuchal, 3966 m; Alamkuh, 4850 m; Damavand, 5671 m), Azarbayjan mountains in NW-Iran (Sahand, 3707 m; Sabalan, 4811 m), and southeastern Zagros (Hezar Mts., 4465 m). In these mountains we were determining the lower limit of the subnival–nival zone by exploring vegetation patterns and species composition. The subnival–nival zone in Central Alborz, where plants still occur, ranges approximately from 3900 to 4800 m, in NW-Iran approximately from 3600 m to 4400 m, and in Hezar–Lalehzar mountains it commences at approximately 4200 m. In Central Zagros the lower limit of the subnival–nival zone is similar to that of Central Alborz. Using the above altitudinal limits, we grouped all species that occur within the subnival–nival zone according to their vertical distribution patterns: Group A comprises the predominantly subnival–nival species, group B subnival–nival to alpine species, and group C those with a wider vertical distribution from subnival to subalpine areas. The grouping is on the species level. Different subspecies, e.g., of Asperula glomerata, range differently through subalpine to nival zones. In these cases we put the species to group C. Some species, such as Arenaria balansae, are restricted to subnival–nival environments in Ira- nian mountains but in other regions such as Anatolia are common in alpine areas. In these 123

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1322 Biodivers Conserv (2011) 20:1319–1338 cases we put these species in group A. Biogeographic relationships of the species of groups A, B and C are calculated separately to understand the relationship between vertical and geographical distributions. Distribution maps of all species of group A (except Luzula spicata) are given. The nomenclature and chorology of species is mainly based on Flora Iranica (Rechinger 1963–2010), and also on Flora of Turkey (Davis 1965–1988), Flora of Caucasus (Gross- heim 1945–1967), Flora of China (http://www.efloras.org/flora_page.aspx?flora_id=2) and the Flora of Pakistan (http://www.efloras.org/flora_page.aspx?flora_id=5). Endemics are defined as species that are restricted to Iran, including cases having their main distribution within the country, but may extend into neighbor countries for a distance of up to c. 30 km.

Results

Altitudinal distribution

A total of 151 vascular plant species, belonging to 27 families and 80 genera, was found to occur within the subnival–nival zone of Iran. This is around 22% of the alpine to nival flora and 2% of the entire flora of Iran. Plant families and genera with the largest number of species and the proportional share of the main life forms of all 151 species are shown in Table 1. The species could be divided into three groups of different vertical distribution patterns (Fig. 2, Table 2). Group A consists of 51 true subnival–nival species, being centred within this uppermost zone, but some species may extend downwards into the upper part of the alpine zone. Group B comprises 56 species being equally distributed across the alpine and the subnival–nival zones. Group C comprises 44 subalpine to alpine species that can extend into subnival areas. Upper climatically determined altitudinal limits of vascular plant life in Iran are reached only in Central Alborz on Damavand (5671 m) and Alamkuh (4850 m), and in the NW of the country on Sabalan (4811 m), the three highest mountain peaks in Iran. In the SE- Zagros range the climatic potential for the upper limit of vascular plant species is estimated to be at around 5000 m, but the highest summit (Hezar mountain) only reaches 4465 m. The altitudinal maximum for flowering plants in Iran according to previous records was

Table 1 The richest plant families and genera and the life forms in the subnival–nival zone of Iran Families No. species Genera No. species Life forms %

Asteraceae 29 Astragalus 8 Hemicryptophyte 70 Caryophyllaceae 15 Potentilla 8 Graminoid 10 Fabaceae 14 Silene 5 Non-cushion chamaephyte 9 Brassicaceae 12 Oxytropis 5 Cushion chamaephyte 4 Lamiaceae 12 Veronica 5 Geophyte 4 Poaceae 11 Draba 4 Annual 3 Scrophulariaceae 9 Nepeta 4 Rosaceae 8 Senecio 4 Apiaceae 6

123

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Fig. 2 Vertical distribution groups of vascular plant species that occur in the subnival–nival zone of Iran, and their percentage endemism to Iran

4800 m (Noroozi et al. 2008). Table 3 shows the species growing in the highest altitudes in Iran and their updated uppermost occurrences. The highest plant community was observed at 4460 m on Damavand; above this elevation, vascular plants only occur as scattered, isolated individuals with low frequency and cover in fragmentary colonies.

Phytogeography

Iran extends over three phytogeographical regions, the Irano–Turanian, Euro–Siberian and the Saharo–Arabian (sensu Zohary 1973) or Sudano–Zambezian (sensu Takhtajan 1986). The alpine area of Iran belongs to the Irano–Turanian region, where the rate of endemism is generally high (Klein 1982, 1991; Noroozi et al. 2008, 2010a, b). Figure 3 illustrates the ﬂoristic relationship of the three altitudinal species groups with mountainous regions outside of Iran and in a global context and Fig. 4 shows the chorological spectra of species with the degree of local endemism. The vertical distribution of species is related to the geographical distribution patterns. The rate of endemism is highest in the true subnival– nival species of group A (68%). Most of the species of group C are widely distributed and the endemism rate is low (20%). Group B takes an intermediate position. Most of endemics are restricted to Central Alborz and Central Zagros. Nine groups of different distribution patterns can be distinguished for the true subnival– nival species (group A): (1) Holarctic distribution: Luzula spicata is the only species being widespread across the northern hemisphere in most alpine and arctic areas (see distribution map in Meusel et al. 1964, p. 88). In Iran this species is only known from Sabalan at 3800 m (Snogerup 1971). Species with similar distribution, but from group B, are Erigeron uniﬂorus (Meusel et al. 1992, p. 457) and Oxyria digyna (Meusel et al. 1964, p. 129). (2) Irano–Turanian to Euro–Siberian distribution: Draba siliquosa (Fig. 5a) and Alope- curus himalaicus (Fig. 5b). Within Iran, both are restricted to very small areas at high elevations. Draba siliquosa reaches its easternmost limit of distribution in Iran and Alopecurus himalaicus has a highly disjunct distribution. Species with similar distribution in group B is Koeleria eriostachya. (3) Hindu Kush–Himalaya system and Central Asian elements with disjunct distribution in Iran: Carex melanantha (Fig. 6a), Kobresia schoenoides (Fig. 6b), K. humilis 123

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Table 2 Species known to occur within the subnival–nival zone of the mountains of Iran Family Species Life form Altitudinal Chorotype group

Alliaceae Allium capitellatum Ge B En(Iran) Apiaceae Carum caucasicum He B Cau-Atro Chaerophyllum nivale He A LEn(Zagos) Chamaesciadium acaule He C Cau-Atro Heracleum anisactis He B LEn(Alboz) Semenovia dichotoma He A En(Zagos) Trachydium depressum He B FI Asteraceae Achillea aucheri NC A En(Atro) Artemisia melanolepis He A En(Atro) Artemisia persica NC C IT Cousinia eburnea He A LEn(Zagros) Cousinia fragilis He A LEn(Zagros) Cousinia lasiolepis NC C FI Crepis heterotricha He B En(Iran) Crepis multicaulis He C Hol Erigeron caucasicus He C Cau-Atro Erigeron hyrcanicus He B En(Alborz) Erigeron uniﬂorus He B Hol Iranecio oligolepis He A LEn(Alborz) Jurinella frigida He B En(Alborz) Myopordon damavandica He A LEn(Alborz) Myopordon persicum He A LEn(Zagros) Psychrogeton alexeenkoi He B IT Psychrogeton amorphoglossus He C IT Psychrogeton chionophilus He A LEn(Zagros) Scorzonera radicosa He B Ana-Atro Senecio iranicus An A LEn(Alborz) Senecio subnivalis An A LEn(Zagros) Senecio taraxacifolius He A Cau-NWIran Senecio vulcanicus He B En(Alborz) Tanacetum kotschyi NC C Ana-Cau-Iran Tanacetum pamiricum NC A IT Taraxacum baltistanicum He A IT Taraxacum crepidiforme He B (Cau)IT Taraxacum primigenium He B En(Zagros) Tripleurospermum caucasicum He C ES(NWIran) Boraginaceae Lepechiniella persica He A En(Alborz) Myosotis asiatica He C Hol Myosotis olympica He B En(Atro) Brassicaceae Aethionema trinervium He C IT Aethionema umbellatum He A LEn(Zagros) Arabis caucasica He C ES-IT Clastopus vestitus He B En(Atro) 123

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Table 2 continued Family Species Life form Altitudinal Chorotype group

Didymophysa aucheri He A Cau-Iran Draba aucheri He B Ir-CAsia Draba bruniifolia He A Ana-Cau-NWIran Draba pulchella He B SEn(Atro) Draba siliquosa He A IT-ES Erysimum caespitosum He C Ana-Iran Erysimum elbrusense He A En(Atro) Graellsia saxifragifolia He C FI Campanulaceae Campanula karakuschensis He A Atro Caryophylaceae Arenaria minutissima NC B LEn(Zagros) Arenaria balansae NC A Ana-Zagros Arenaria insignis CC C FI Cerastium cerastioides He C Hol Cerastium purpurascens He A Ana-Cau-Atro Dianthus erythrocoleus NC B Atro Herniaria caucasica He C Ana-Cau-Atro Minuartia aizoides CC B NAna-Cau-NWIran Minuartia lineata CC C SEn(Atro) Minuartia recurva He C ES-IT Silene dae¨nensis He A En(Zagros) Silene nurensis CC B En(Zagros) Silene odontopetala He C IT Silene persica He B En(Zagros) Silene tragacantha He A LEn(Zagros) Chenopodiaceae Chenopodium foliosum An C PL Crassulaceae Sedum tenellum An B Cau-Atro Sedum kotschyanum He B En(Zagros) Cyperaceae Carex melanantha Gr A IT Kobresia humilis Gr A IT Kobresia schoenoides Gr A (Cau)IT Euphorbiaceae Euphorbia aucheri He C FI Euphorbia cheiradenia He C En(Iran) Fabaceae Astragalus atricapillus He A En(Alborz) Astragalus capito He B En(Alborz) Astragalus macrosemius CC B En(Alborz) Astragalus melanocalyx He A LEn(Zagros) Astragalus melanodon He C En(Zagros) Astragalus montis-varvashti He B LEn(Alborz) Astragalus tenuiscapus He C En(Zagros) Astragalus zerdanus He B En(Zagros) Cicer stapﬁanum He A LEn(Zagros)

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Table 2 continued Family Species Life form Altitudinal Chorotype group

Oxytropis immersa He B IT Oxytropis kermanica He A En(Iran) Oxytropis persica He B SEn(Atro) Oxytropis savellanica He B IT Oxytropis takhti-soleimanii He A LEn(Alborz) Gentianaceae Gentiana pontica He B EAna-Cau-Alborz Juncaceae Luzula spicata Gr A Hol Lamiaceae Betonica nivea He B ES-IT Dracocephalum aucheri NC A SEn(Atro) Dracocephalum surmandinum NC A LEn(Zagros) Lamium tomentosum He A Cau-Atro Nepeta allotria He B LEn(Alborz) Nepeta archibaldii He A LEn(Zagros) Nepeta crispa He B En(Iran) Nepeta lasiocephala He B LEn(Zagros) Scutellaria glechomoides He A LEn(Alborz) Stachys obtusicrena He B En(Zagros) Thymus pubescens NC C Atro Ziziphora clinopodioides NC C IT Liliaceae Gagea alexii Ge A FI Gagea caroli-kochi Ge B Atro Gagea soleimani Ge C En(Atro) Plantaginaceae Plantago atrata He C ES-IT Plumbaginaceae Acantholimon demawendicum CC B En(Alborz) Acantholimun haesarense CC A LEn(Zagros) Poaceae Alopecurus dasyanthus Gr B Ana-Cau-NWIran Alopecurus himalaicus Gr A ES-IT Alopecurus textilis Gr C IT Bromus frigidus Gr A En(Zagros) Bromus tomentosus Gr C (Cau)IT Catabrosella parviﬂora Gr C (Cau)IT Festuca alaica Gr C FI Koeleria eriostachya Gr B ES(NWIran) Poa alpina Gr B Hol Poa araratica Gr C (Cau)IT Stipa hohenackeriana Gr C IT Polygonaceae Oxyria digyna He B Hol Polygonum serpyllaceum He B FI Primulaceae Androsace villosa He B ES-IT Primula algida He B (Cau)IT

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Table 2 continued Family Species Life form Altitudinal Chorotype group

Ranunculaceae Paraquilegia caespitosa He A En(Alborz) Ranunculus crymophilus He B SEn(Atro) Ranunculus eriorrhizus He B En(Zagros) Rosaceae Potentilla argaea He B Ana-Cau-NWIran Potentilla argyroloma He B En(Iran) Potentilla aucheriana He B SEn(Atro) Potentilla gelida He B ES-IT Potentilla hololeuca He B Iran-CAsia Potentilla nuda He C En(Iran) Potentilla polyschista He A SEn(Atro) Potentilla porphyrantha He A SEn(Atro) Rubiaceae Asperula glomerata NC C (Cau)IT Galium aucheri He A En(Alborz) Galium decumbens He B En(Atro) Saxifragaceae Saxifraga exarata He C ES-IT Saxifraga iranica He A LEn(Alborz) Saxifraga sibirica He C ES-IT Scrophulariaceae Euphrasia juzepczukii An C Atro Pedicularis caucasica He B Cau-Atro Scrophularia olympica He B Ana-Cau-NWIran Scrophularia subaphylla He C En(Iran) Veronica aucheri He A En(Alborz) Veronica biloba An C IT Veronica gentianoides He C Ana-Cau-Atro Veronica kurdica He C En(Iran) Veronica paederotae He A LEn(Alborz) Woodsiaceae Cystopteris fragilis He C PL Life forms: An annual, Ge geophyte, CC cushion chamaephyte, NC non-cushion chamaephyte, He Hemi- cryptophyte, Gr graminoid. Chorology: Hol Holarctic, PL Pluriregional, IT Irano–Turanian, ES Euro– Siberian, Atro Atropatenian subprovince, FI Flora Iranica territory, Ana Anatolia, Cau Caucasus, En endemic, LEn local endemic, SEn subendemic. Altitudinal groups: A true subnival-nival, B alpine–subnival, C subalpine–subnival

(Fig. 6c), Taraxacum baltistanicum (Fig. 6d), Tanacetum pamiricum (Fig. 6e) and Gagea alexii (Fig. 6f). Species with similar distribution in group B are Psychrogeton alexeenkoi (see distribution map in Noroozi et al. 2010a), Draba aucheri, Oxytropis immersa, O. savellanica and Potentilla hololeuca. The distribution of several genera or species groups within genera ﬁt to this pattern. In the genus Paraquilegia, consisting of 12 species, 11 species are found in the Hindu Kush, Himalaya and in Central to Middle Asia and one, P. caespitosa, is endemic to Iran and grows in subnival–nival rocky habitats of Central Alborz, forming the westernmost outpost of the genus. The Iran-endemic Dracocephalum aucheri is a member of a distinctive species group, containing several species that stretch

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Table 3 The uppermost records of vascular plants on the four highest mountains of Iran Central Alborz NW-Iran SE-Zagros

Damavand (5671 m) Alamkuh (4850 m) Sabalan (4811 m) Hezar (4465 m)

4820 m 4735 m 4371 m 4450 m Veronica aucheri Saxifraga iranica Draba bruniifolia Artemisia persica 4799 m 4730 m Potentilla porphyrantha Asperula glomerata Didymophysa aucheri Cerastium 4357 m Astragalus melanodon Achillea aucheri purpurascens Alopecurus dasyanthus Gagea alexii Alopecurus himalaicus Veronica aucheri Didymophysa aucheri Nepeta lasiocephala 4767 m Draba siliquosa Potentilla nuda Erysimum elbrusense Erigeron caucasicus Psychrogeton 4600 m Saxifraga sibirica alexeenkoi Dracocephalum Tripleurospermum P. amorphoglosus aucheri caucasicum Ranunculus eriorrhizus Scrophularia subaphylla Senecio subnivalis Silene dae¨nensis

Fig. 3 Global distribution of vascular plants occurring in the subnival–nival zone of Iran: percentages of Iranian endemic species and of other regions where wider-spread species occur; compare text and Fig. 2 for species groups A (a), B (b) and C (c)

Fig. 4 Chorological spectra of species in groups A (a), B (b) and C (c). IT(Zag) endemic of Zagros, IT(Alb) endemic of Alborz, IT(Ir) endemic of Iran, IT Irano–Turanian, IT-ES: Irano–Turanian/Euro–Siberian, PL: Pluriregional

disjunctly from SE Turkey to the Alborz, E Afghanistan, C Asia and thence to the extreme NE of Russia (Hedge and Wendelbo 1978). Saxifraga iranica (also endemic to Iran) is very close to two Himalayan species, S. imbricata and S. namulosa (Bornmu¨ller 1906 in Klein 1991), belonging to the Marginatae group with 19 species, 123

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Fig. 5 Irano–Turanian to Euro–Siberian distribution: a Draba siliquosa and b Alopecurus himalaicus

where 13 species are distributed in Himalaya and China (Engler and Irmscher 1919 in Klein 1991). (4) Distributed from Iran to Caucasus and/or Anatolia: Draba bruniifolia (Fig. 7a), Cerastium purpurascens (Fig. 7b) and Arenaria balansae (Fig. 7c). Species with similar distribution in group B are Scrophularia olympica, Scorzonera radicosa (Meusel et al. 1992, p. 532) and Potentilla argaea. (5) Distributed from Iran to the Caucasus and E Anatolia: Didymophysa aucheri (Fig. 8a), Lamium tomentosum (Fig. 8b), Senecio taraxacifolius (Fig. 8c) and Campanula karakuschensis (Fig. 8d). Species with similar distribution in group B are Carum caucasicum, Gentiana pontica (Meusel et al. 1964, p. 353), Minuartia aizoides and Pedicularis caucasica (Noroozi et al. 2008). (6) Distributed across the Atropatenian subprovince sensu Takhtajan (1986): Draco- cephalum aucheri (Fig. 9a), Potentilla polyschista (Fig. 9b), P. porphyrantha (Fig. 9c), Achillea aucheri (Fig. 9d), Artemisia melanolepis (Fig. 9e) and Erysimum elbrusense (Fig. 9f). The Atropatenian subprovince includes the arid and semiarid parts of Transcaucasia, the east and southeast of Turkey, the southern Armenian highlands, the northwest of Iran, and the Alborz mountains. This subprovince is one of the most active centers of speciation in Western Asia (Takhtajan 1986). Species with similar distribution in group B are Myosotis olympica, Dianthus erythrocoleus, Clastopus vestitus (see distribution map in Hedge and Wendelbo 123

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Fig. 6 Hindukush–Himalaya system and Central Asia elements with disjunct distribution. a Carex melanantha, b Kobresia schoenoides, c K. humilis, d Taraxacum baltistanicum, e Tanacetum pamiricum, f Gagea alexii

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Fig. 7 Distributed from Iran to Caucasus and/or Anatolia. a Draba bruniifolia; subsp. bruniifolia (circle), other subspecies (triangular), b Cerastium purpurascens, c Arenaria balansae

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Fig. 8 Distributed from Iran to the Caucasus and E Anatolia. a Didymophysa aucheri, b Lamium tomentosum, c Senecio taraxacifolius and d Campanula karakuschensis

1978), Draba pulchella (Noroozi et al. 2008), Oxytropis persica (Noroozi et al. 2008), Ranunculus crymophilus, Potentilla aucheriana (Noroozi et al. 2008) and Gagea caroli-kochii. (7) Distributed in Alborz and Zagros: Oxytropis kermanica (Fig. 9g). Species with similar distribution in group B are Potentilla argyroloma, P. nuda (Noroozi et al. 2010a), Scrophularia subaphylla (Noroozi et al. 2010a), and Veronica kurdica (Noroozi et al. 2008). (8) Alborz endemic species: Galium aucheri (Fig. 10a), Lepechinella persica (Fig. 10b), Astragalus atricapillus (Fig. 10c), Veronica aucheri (Fig. 10d), V. paederotae (Fig. 10e), Paraquilegia caespitosa (Fig. 10f), Saxifraga iranica (Fig. 10g), Iranecio oligolepis (Fig. 10g), Scutellaria glechomoides (Fig. 10h), Myopordon damavandica (Fig. 10h), Oxytropis takhti-solimanii (Fig. 10i) and Senecio iranicus (Fig. 10i). Species with similar distribution in group B are Erigeron hyrcanicus (Meusel et al. 1992, p. 457), Senecio vulcanicus (Noroozi et al. 2008), Jurinella frigida, Astragalus capito, A. macrosemius, A. montis-varvashti, Nepeta allotria, Acantholimon demawendicum (Noroozi et al. 2008), Heracleum anisactis, Galium decumbens, Erigeron uniﬂorus subsp. elbursensis and Asperula glomerata subsp. bracteata.

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Fig. 9 Distributed across the Atropatenian subprovince. a Dracocephalum aucheri, b Potentilla polyschista, c P. porphyrantha, d Achillea aucheri [subsp. aucheri (circle), subsp. glabra (triangle)], e Artemisia melanolepis, f Erysimum elbrusense and distributed in Alborz and Zagros. g Oxytropis kermanica

(9) Zagros endemic species: Bromus frigidus (Fig. 11a), Chaerophyllum nivale (Fig. 11a), Acantholimon haesarense (Fig. 11a), Aethionema umbellatum (Fig. 11b), Cousinia eburnea (Fig. 11b), C. fragilis (Fig. 11b), Myopordon persicum (Fig. 11c), Dracocephalum surmandicum (Fig. 11c), Nepeta archibaldii (Fig. 11d), Silene tragacantha (Fig. 11d), Semenovia dichotoma (Fig. 11e), Silene dae¨nensis (Fig. 11f), Astragalus melanocalyx (Fig. 11f), Psychrogeton chionophilus (Fig. 11g), Cicer stapﬁanum (Fig. 11g), Senecio subnivalis (Fig. 11g) and Asperula glomerata subsp. condensata (Fig. 11h). Species with similar distribution in group B are Arenaria minutissima, Astragalus zerdanus, Erigeron uniﬂorus subsp. dae¨nensis, Nepeta lasiocephala (Noroozi et al. 2010a), Ranunculus eriorrhizus (Noroozi et al. 2010a), Silene nurensis (Noroozi et al. 2010a), S. persica, Stachys obtusicrena (Noroozi et al. 2008) and Taraxacum primigenium (Noroozi et al. 2010a). The degree of endemism appears to be higher in Zagros mountains compared with Alborz (Wendelbo 1971; Akhani 2004, 2007; Noroozi et al. 2008, 2010a).

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Fig. 10 Alborz endemic species. a Galium aucheri, b Lepechinella persica, c Astragalus atricapillus, d Veronica aucheri, e V. paederotae, f Paraquilegia caespitosa, g Saxifraga iranica (circle), Iranecio oligolepis (triangle), h Scutellaria glechomoides (circle), Myopordon damavandica (triangle), i Oxytropis takhti-solimanii (circle) and Senecio iranicus (triangle)

Discussion

Endemism, species vertical ranges and altitude

The rate of endemism increases very sharply with increasing altitude in Iran, where a narrow vertical distribution restricted to high altitudes often corresponds to constricted geographic range. Vice versa, species with a larger amplitude of their altitudinal distribution commonly are widespread species (see Figs. 2, 3 and 4). Similar patterns were observed in several European mountains, for example in mountains exposed to Mediter- ranean-type climate such as the Sierra Nevada (Spain) (Pauli et al. 2003) and the central Apennines (Italy) (Stanisci et al. 2005), but also in some mountain ranges in the temperate climate of Central Europe. For example, in the NE Calcareous Alps the highest rate of endemism was observed in the upper elevation zones (subalpine to alpine) of these lower and outer parts of the Alps, whereas the percentage of endemism generally decreases towards the central Alps; the latter having been fully glaciated during the Pleistocene (Pauli et al. 2003; Essl et al. 2009). Also in the Caucasus mountains the percentage of endemics was reported to be highest in the subnival belt (Nakhutsrishvili 1998). In the Hindu Kush mountains, however, the percentage of endemism peaks in the subalpine zone, but species dwelling in the nival zone (above 5000 m) mostly have a wide geographical distribution over the Eurasian mountains and the Arctic (Breckle 1974, 2004). Similar 123

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Fig. 11 Zagros endemic species. a Bromus frigidus (circle), Chaerophyllum nivale (triangle), Acanthol- imon haesarense (square), b Aethionema umbellatum (circle), Cousinia eburnea (triangle), C. fragilis (square), c Myopordon persicum (triangle), Dracocephalum surmandicum (square), d Nepeta archibaldii (circle), Silene tragacantha (triangle), e Semenovia dicotoma, f Silene dae¨nensis (circle), Astragalus melanocalyx (triangle), g Psychrogeton chionophilus (triangle), Cicer stapfianum (circle) and Senecio subnivalis (square), h Asperula glomerata subsp. condensata distribution patterns were found in the Himalaya (Vetaas and Grytnes 2002). This contrast may be explained by the lesser orographic isolation of the Hindu Kush and Himalaya in the past and present, compared to the high mountains of Iran. Orogenesis during the Cenozoic played a major role in the evolution of the alpine flora (Agakhanjanz and Breckle 1995). The high rate and extent of mountain uplift in the Alborz and Zagros and the high degree of fragmentation and isolation of their high-elevation areas from the neighboring mountain ranges (Breckle 2009) are considered as important determinants for the pronounced high-elevation endemism (Noroozi et al. 2008, 2010a). In spite of extensive glaciations in the mountains of Iran and a climatic snowline level lowered by 600–1,100 m during the Pleistocene (Bobek 1937; Ferrigno 1991), the glaciations may have not severely affected the overall distribution patterns of the flora of Alborz and Zagros mountains (Bobek 1937, 1953; Noroozi et al. 2008). Mainly valley glaciations may have occurred (Agakhanjanz and Breckle 1995), thus enabling many high-mountain species to persist on relict stands not far from their current occurrences (Hedge and Wendelbo 1978; Klein 1991; Noroozi et al. 2008, 2010a). A strong orographic isolation and the absence of larger ice-shields during the ice-ages appear to be crucial for the present high- mountain endemism in Iran and other mountain regions where a similar situation occurs, such as the Spanish Sierra Nevada (Blanca et al. 1998) or the Moroccan High Atlas. 123

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A general south–north gradient of decreasing endemism from the Atlas mountains to northern Europe can be found, which coincides with an increasing Pleistocene glaciation and connectivity among cold areas (Favarger 1972). The surprisingly low degree of alpine endemic plants in North America (Mills and Schwartz 2005) may be associated with maintained connections and migration pathways supported by extensive north–south-oriented mountain ranges. Climatic links during the Pleistocene glacial periods between the Himalaya and Central Asia with the Iranian, Caucasian and some European mountains facilitated a long-distance migration of several cryophilic species. Through the post-glacial climatic amelioration and, thus, the disruption of migration corridors, the distribution of some species became highly disjunct (e.g., Alopecurus himalaicus, Fig. 5b). In addition to a high level of endemism, the pronounced orographic isolation in Iran led to an abrupt decrease of plant diversity with altitude (Noroozi et al. 2008). By contrast, the past and contemporary connections of cold areas of Hindu Kush to neighboring ranges (Himalaya, Central and Middle Asia) facilitated species migrations (Breckle 2007).

Global warming and the vulnerability of subnival–nival endemics

In spite of the importance of historic determinants, the temperature component of orographic isolation is the most important factor favouring endemism in high mountain areas at present (Pauli et al. 2003). A changing temperature regime through an ongoing climate warming therefore may increasingly threaten the persistence of the isolated and unique cryophilic ﬂora of Iran, where the potential to escape to suitable cold habitats is highly limited. The narrower the altitudinal distribution, the geographical range and the within- species genetic diversity and plasticity, the more severe is the potential vulnerability to climate change (Dirnbo¨ck et al. 2003; Ohlemu¨ller et al. 2008). This particularly accounts for the endemic species of group A and, to a lesser extent, of group B of the Iranian subnival–nival ﬂora. There is evidence of an ongoing upward migration (e.g., Grabherr et al. 1994; Walther et al. 2005) and of a decline of subnival–nival species at their lower range margins in the Alps (Pauli et al. 2007). No such data are available for the Iranian mountains owing to the lack of permanent observation plots. First sites in the Alborz mountains only were setup in 2008 by the authors as part of the GLORIA network (www.gloria.ac.at). Unlike the situation in the Alps, but similar to other Mediterranean-type mountains, barriers against invading competitor species from lower elevations, such as closed forest and grassland belts, are absent in Iranian mountains. Instead, open habitats from lower elevations up to the high terrains may support an upward colonization driven by climate change. Hypothetically, climate warming is coupled with an earlier snow-melt and, thus, to an extended low-precipitation period in summer that, in turn, may lead to a reduced vitality of cryophilic plants. A prolonged drought period could also affect invaders from lower elevations, but these species are likely to be well adapted to arid conditions (Pauli et al. 2003).

Conclusions

(1) In Iran, 151 vascular plant species were identiﬁed to occur in the subnival–nival zone, where 51 of these can be considered as true subnival–nival species. Within the latter,

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the degree of species endemic to Iran is 68% and clearly decreases to 53 and 20% for species that also occur in the alpine and in the subalpine zones, respectively. (2) The outstanding level of high-altitude endemism appear to result mainly from a pronounced orographic isolation of the country’s highly scattered and fragmented cold areas and the absence of extensive Pleistocene glaciations. (3) Predicted climate warming may seriously threaten the survival of the unique subnival– nival ﬂora of Iran, due to a very low potential of alternative low-temperature habitats at higher elevations. Additional pressure through drought-tolerant competitors from lower altitudes can further increase the risk of biodiversity losses. (4) An expansion of long-term observation sites, and of ecological, biogeographic and phylogeographic research on Iran’s high-altitude ﬂora and vegetation is strongly recommended.

Acknowledgments Parts of traveling and subsistence costs for this study were funded by the GLORIA coordination (University of Vienna and Austrian Academy of Sciences/Institute of Mountain research), the association GLORIA-International, Vienna and the Austrian Federal Ministry of Science and Research. We would like to thank Yousef Ajani for his assistance during ﬁeldwork and Iris Wagner for her help in drawing the distribution maps.

References

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Essl F, Staudinger M, Sto¨hr O et al (2009) Distribution patterns, range size and niche breadth of Austrian endemic plants. Biol Conserv 142:2547–2558 Favarger C (1972) Endemism in the montane floras of Europe. In: Valentine DH (ed) Taxonomy, phytogeography and evolution. Academic Press, London, pp 191–204 Ferrigno JG (1991) Glaciers of the Middle East and Africa—glaciers of Iran. In: Williams RS, Ferrigno JG (eds) Satellite image atlas of glaciers of the world. U.S. Geological Survey Professional Paper 1386-G. United States Government Printing Office, Washington, pp 31–47 Gilli A (1939) Die Pflanzengesellschaften der Hochregion des Elbursgebirges in Nordiran. Beih Bot Cbl 59:317–344 Gilli A (1941) Ein Beitrag zur Flora des Elburs-Gebirges in Nord-Iran. Feddes Repert Spec Regni Veg 50:263–283 Grabherr G, Gottfried M, Pauli H (1994) Climate effects on mountain plants. Nature 369:448 Grossheim AA (1945–1967) Flora Kavkaza (Flora of Caucasus), vols 3–7. Nauk, Leningrad Hedge IC, Wendelbo P (1978) Patterns of distribution and endemism in Iran. Notes Roy Bot Gard Edinb 36:441–464 Klein JC (1982) Les groupements chionophiles de l’Alborz central (Iran). Comparaison avec leurs homolegues d’ Asie centrale. Phytocoenologia 10(4):463–486 Klein JC (1991) Endemisme a` l’etage alpin de l’Alborz (Iran). Flora Veg Mundi 9:247–261 Klein JC, Lacoste A (1999) Observation sur la ve´ge´tation des e´boulis dans les massifs irano-touraniens: le Galietum aucheri ass. nov. de l’Alborz Central (N-Iran). Documents phytosociologiques 19:219–228 Kotschy T (1861a) Die Vegetation des westlichen Elbrus in Nordpersien. O¨ sterr Bot Zeitschr 11(4):105–117 Kotschy T (1861b) Der westliche Elbrus bei Teheran. Mitt. der K. K. Oester Geogr Gesell Wien 5:65–110 Meusel H, Ja¨ger E, Weinert E (1964) Vergleichende Chorologie der zentraleuropaïschen Flora Band I. Fischer, Jena Meusel H, Ja¨ger E, Braütigam S et al (1992) Vergleichende Chorologie der zentraleuropaïschen Flora, Band III. Fischer, Jena Mills MH, Schwartz MW (2005) Rare plants at the extremes of distribution: broadly and narrowly distributed rare species. Biodivers Conserv 14:1401–1420 Nagy L, Grabherr G (2009) The biology of alpine habitats. Oxford University Press, New York Nakhutsrishvili G (1998) The vegetation of the subnival belt of the Caucasus Mountains. Arct Alp Res 30:222–226 Noroozi J, Akhani H, Breckle S-W (2008) Biodiversity and phytogeography of the Alpine flora of Iran. Biodivers Conserv 17:493–521 Noroozi J, Ajani Y, Nordenstam B (2010a) A new annual species of Senecio (Compositae–Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area. Comp Newsl 48:4–23 Noroozi J, Akhani H, Willner W (2010b) Phytosociological and ecological study of the high alpine vegetation of Tuchal Mountains (Central Alborz, Iran). Phytocoenologia 40(4):293–321 Ohlemu¨ller R, Anderson BJ, Araujo MB et al (2008) The coincidence of climatic and species rarity: high risk to small-range species from climate change. Biol Let 4:568–572 Pauli H, Gottfried M, Dirnbo¨ck T et al (2003) Assessing the long-term dynamics of endemic plants at summit habitats. In: Nagy L, Grabherr G, Ko¨rner C et al (eds) Alpine biodiversity in Europe—a Europe-wide assessment of biological richness and change, Ecological Studies. Springer, Berlin, vol 167, pp 195–207 Pauli H, Gottfried M, Reiter K et al (2007) Signals of range expansions and contractions of vascular plants in the high Alps: observations (1994–2004) at the GLORIA master site Schrankogel, Tyrol, Austria. Glob Change Biol 13:147–156 Rechinger KH (ed) (1963–2010) Flora Iranica, vols 1–178. Akad. Druck-u. Verlagsanstalt, Graz Snogerup S (1971) Juncaceae. In: Rechinger KH (ed) Flora Iranica, No. 75. Akademische Druck u. Ver- lagsanstalt, Graz Stanisci A, Pelino G, Blasi C (2005) Vascular plant diversity and climate change in the alpine belt of the central Apennines (Italy). Biodivers Conser 14:1301–1318 Takhtajan A (1986) Floristic regions of the world. University of California Press, California (English translation from Russian) Vetaas OR, Grytnes JA (2002) Distribution of vascular plant species richness and endemic richness along the Himalayan elevation gradient in Nepal. Glob Ecol Biogeogr 11:291–301 Walther GR, Beißner S, Burga CA (2005) Trends in upward shift of alpine plants. J Veg Sci 16:541–548 Wendelbo P (1971) Some distributional patterns within the Flora Iranica area. In: Davis PH, Harper PC, Hedge IC (eds) Plant life of South–West Asia. Edinburgh, pp 29–41 Zohary M (1973) Geobotanical foundations of the Middle East, vol 2. Fischer, Stuttgart

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Title of manuscript: High Mountain Regions in Iran

Author: Jalil Noroozi

Status: in press, (chapter 7.6), in C. Hobohm (editor). Endemism in Vascular Plants. Springer.

Contribution: idea, data collection, data analysis, manuscript writing.

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7.6 High Mountain Regions in Iran Jalil Noroozi

7.6.1 Physical Geography of Mountain Ranges in Iran

Iran with a total surface area of c. 1.6 million km² is a typical high mountain country. Almost half the country consists of high elevations, and Alborz and Zagros are the major high mountain chains (Fig. 7.7). The highest mountains are Damavand (5671 m asl.), Alamkuh (4850 m), Sabalan (4810 m), and Hezar (4465 m). More than 100 mountain peaks exceed 4000 m. There are glaciers in the higher elevations, i.e. Damavand, Alamkuh, Sabalan and Zardkuh (Ferrigno 1991). According to Schweizer (1972) the present snowline in Alborz, north and central Zagros, and the NW Iranian mountains lies between 4000 and 4200 m. In the mountains of central and southern Iran, e.g. Shirkuh, Dena, and Hezar-Lalehzar Mts. it is between 4500 to 5000 m. The higher elevations of Iran have a continental climate with Mediterranean precipitation regime. The annual precipitation in the higher altitudes of Alborz reaches almost 1000 mm (Khalili 1973). The alpine habitats are almost all above 3000 m, and are found scattered across different parts of the country (see Fig. 7.7).The Iranian alpine flora is of Irano-Turanian origin (Zohary 1973, Klein 1982, 1991, Frey et al. 1999). A conspicuous feature of this flora is the high rates of endemism. A total of 682 vascular plant species are known from alpine habitats of Iranian mountains, and of this total 394 species are endemic or subendemic to Iran (Noroozi et al. 2008). These habitats are less known, and plant species are still being discovered and described as new to science (e.g. Jamzad 2006, Khassanov et al. 2006, Noroozi et al. 2010a, Noroozi & Ajani 2013, Razyfard et al. 2011). Several different vegetation types are found in the high regions of Iran. The dry slopes of the subalpine zone are usually covered with tall herbs and umbelliferous vegetation types (Klein 1987, 1988). Small patches of subalpine wetlands are found scattered on dry slopes (Naqinezhad et al. 2010). Thorn-cushion grasslands occur in alpine meadow and windswept areas, and snowbed vegetation types in snow patches or

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snow melting runnels (Klein 1982, Noroozi et al. 2010b). Scree vegetation types with sparse vegetation cover occupy high alpine and subnival steep slopes with a high percentage of screes and stones on the ground (Klein & Lacoste 2001, Noroozi et al. 2013).

Fig. 7.7 Alborz and Zagros are the major mountain chains of Iran. The alpine habitats (black spots) are scattered around different parts of the country.

7.6.2 Analysis of Floras and Phytosociological Investigation

To select the local endemic species, the Flora Iranica (Rechinger 1963-2012) and Flora of Iran (Assadi et al. 1988-2012) were used as the main sources.

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The main data on endemism in alpine habitats of Iran and the percentage of endemism in alpine and subnival-nival zones were extracted from Klein (1991), Hedge & Wendelbo (1978) and Noroozi et al. (2008, 2011). The rate of endemism in different habitats is based on phytosociological vegetation studies of the author in Central Alborz. This rate is measured from all character species which were recorded for each habitat. Some species which were characteristic for two different habitats were separately calculated for both habitats. For example Astragalus macrosemius is a character species for alpine thorn-cushion grasslands and alpine- subnival scree grounds and was thus counted separately for both habitats as a character species.

7.6.3 Endemism in Different High Mountain Areas and Habitats

Three monotypic genera are found exclusively within Iranian alpine habitats, Elburzia in Central Alborz (Hedge 1969), Sclerochorton in northern Zagros (Rechinger 1987), and Zerdana in central, south and southeastern Zagros (Rechinger 1968). There are also some interesting ditypic genera in these habitats, such as Clastopus with two species endemic to Iran (Hedge & Wendelbo 1978), Didymophysa with one species occurring in Iran and Transcaucasus, and another one in central Asia and Hindukush (Hedge 1968), and Dielsiocharis with one endemic species in Iran (Hedge 1968) and one local endemic in central Asia (Al-Shehbaz & Junussov 2003). All the above-mentioned genera belong to the Brassicaceae except Sclerochorton, which belongs to Apiaceae. The author has only seen Zerdana, Clastopus and Didymophysa in the wild, and all of these are restricted to scree habitats. A total of 110 vascular plant species were considered to be rare and narrow endemic to Iranian alpine habitats and have only been recorded from one or very few locations. These species are found in Zagros, Alborz, and in the northwestern part of the country (Sahand and Sabalan Mts.). They are classified into four mountainous regions as below: Rare species in Alborz: Alchemilla amardica, A. rechingeri, Allium tuchalense, Astragalus aestivorum, A. herbertii, A. montis-varvashti, A. nezva-montis, Cousinia decumbens, C. harazensis,

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Diplotaenia damavandica, Elburzia fenestrata, Erodium dimorphum, Festuca rechingeri, Galium delicatulum, Iranecio oligolepis, Myopordon damavandica, M. hyrcanum, Nepeta allotria, N. pogonosperma, Oxytropis aellenii, O. cinerea, O. takhti-soleimanii, Paraquilegia caespitosa, Phlomis ghilanensis, Saxifraga koelzii, S. ramsarica, S. iranica, Scorzonera xylobasia, Scutellaria glechomoides, Senecio iranicus, Silene demawendica, Thlaspi maassoumi, Trachydium eriocarpum, Veronica euphrasiifolia, V. paederotae, Vicia aucheri. Rare species in the mountains of northwestern Iran: Astragalus azizii, A. pauperiflorus, A. savellanicus, Dianthus seidlitzii, Euphorbia sahendi, Festuca sabalanica, Nepeta sahandica, Ranunculus renzii, R. sabalanicus, Thlaspi tenue. Rare species in north, central and southern Zagros (from Kordestan to Fars provinces): Acantholimon eschkerense, Allium mahneshanense, Arenaria minutissima, Astragalus mahneshanensis, A. montis-parrowii, Bufonia micrantha, Chaerophyllum nivale, Cicer stapfianum, Cousinia concinna, C. eburnea, Crepis connexa, Cyclotrichium straussii, Dianthus elymaiticus, Dionysia aubrietioides, D. iranshahrii, Dracocephalum surmandinum, Erysimum frigidum, Euphorbia plebeia, Festuca iranica, Galium schoenbeck-temesyae, Jurinea viciosoi, Myopordon aucheri, Nepeta archibaldii, N. chionophila, N. iranshahrii, N. monocephala, N. natanzensis, Potentilla flaccida, Psychrogeton chionophilus, Ranunculus dalechanensis, Salvia lachnocalyx, Satureja kallarica, Sclerochorton haussknechtii, Scorzonera nivalis, Scrophularia flava, Senecio kotschyanus, Seratula melanocheila, Silene hirticalyx, Tragopogon erostris, Veronica daranica. Rare species in southeastern Zagros (mountains of Yazd, Kerman and Baluchestan provinces): Acantholimon haesarense, A. kermanense, A. nigricans, A. sirchense, Allium lalesaricum, Astragalus hezarensis, A. melanocalyx, A. pseudojohannis, Chaenorhinum grossecostatum, Cousinia fragilis, Dionysia curviflora, Helichrysum davisianum, Hymenocrater yazdianus, Hyoscyamus malekianus, Nepeta asterotricha, N. bornmulleri, N. rivularis, Polygonum spinosum, Rubia caramanica, Senecio eligulatus, S. subnivalis, Silene dschuparensis, Verbascum carmanicumm.

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Since alpine habitats have been less well investigated than lower elevations, our knowledge about the distribution range of species living in high mountain regions is low. This means that it is more likely that new explorations at these elevations will produce new records and localities for local endemics. Nonetheless, the above mentioned species are rare, with narrow geographical and altitudinal distribution. Most of the above- mentioned species can be categorised as Endangered (EN) and Critically Endangered (CR) according to IUCN Red List criteria. However, more field studies are needed to clarify this. Some of these species are only known from type specimens. For instance, Sclerochorton haussknechtii has not been found for 140 years, since it was first collected. This could be because the species has become extinct in the wild or, more likely, because of a lack of information due to a scarcity of field investigations. Based on the vegetation data of the author from Central Alborz, the highest rate of endemism occurs in alpine-subnival scree grounds (60% taxa), followed by alpine thorn- cushion grasslands (49% taxa), subalpine dry slopes covered with umbelliferous vegetation types (41% taxa), alpine snowbeds (36% taxa) and subalpine wetlands (6% taxa). Examples for endemics adapted to alpine-subnival scree grounds are: Asperula glomerata subsp. bracteata, Astragalus capito, Cicer tragacanthoides, Clastopus vestitus, Crepis heterotricha, Dracocephalum aucheri, Galium aucheri, Jurinella frigida, Leonurus cardiaca ssp. persicus, Nepeta racemosa, Scutellaria glechomoides, Senecio vulcanicus, Veronica paederotae, Veronica aucheri and Ziziphora clinopodioides subsp. elbursensis. Examples for endemics which occur in alpine thorn-cushion grasslands are: Acantholimon demawendicum, Allium tuchalense, Astragalus chrysanthus, A. iodotropis, A. macrosemius (Photo 7.14), Bufonia kotschyana, Cousinia crispa, Draba pulchella, Minuartia lineata, Oxytropis persicus, Scorzonera meyeri, Silene marschallii, Tragopogon kotschyi and Veronica kurdica.

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Photo 7.14 Astragalus macrosemius (3700 m) in Central Alborz with Mt. Damavand (5671 m) in the background (photographed by Noroozi)

Examples of endemics which inhabit subalpine dry slopes covered with umbelliferous vegetation are: Aethionema stenopterum, Allium derderianum, A. elburzense, Astragalus aegobromus, Cousinia adenosticta, C. hypoleuca, Echinops elbursensis, Galium megalanthum, Iris barnumae, Parlatoria rostrata, Ranunculus elbursensis and Rumex elbursensis. Examples of endemic taxa of alpine snowbeds are: Cerastium persicum, Erigeron uniflorus ssp. elbursensis, Potentilla aucheriana and Ranunculus crymophylus. An example of endemics of subalpine wetlands is Ligularia persica. According to Naqinezhad et al. (2010), of 323 vascular plant species recorded for the wetland flora on the southern slopes of Alborz, only 7% are endemic and subendemic to Iran, which is consistent with our local findings. This rate of endemism is very low in comparison to other high-elevation habitats in Iranian mountains. The percentage of endemic species for the true subnival-nival flora is 68% (Noroozi et al. 2011), and for alpine habitats (including the subnival-nival zone) 58% (Noroozi et

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al. 2008). This rate is much higher than that for the entire Iranian flora (24%, Akhani 2006). It means that c. 23% of Iranian endemics are concentrated in alpine habitats, suggesting that the alpine zone can be considered one of the centres of endemism. Most studies on plant biodiversity of Iranian mountains demonstrate that the proportion of narrowly distributed plant species increases consistently from low to high elevations (Klein 1991, Noroozi et al. 2008, 2011, Naqinezhad et al. 2009, 2010, Kamrani et al. 2011). The major factors increasing the extinction risk for geographically restricted alpine species of Iran could be climate change and overgrazing. The evidence of global warming impacts on the upward shift of plant species has been demonstrated for European mountains (e.g., Grabherr et al. 1994, Walther et al. 2005, Gottfried et al. 2012, Pauli et al. 2012). The persistence of the unique cryophilic flora of Iran would be seriously threatened under the impact of ongoing global warming where the potential to migrate to appropriate habitats is very limited (Noroozi et al. 2011). High elevations in Iran have been used as summer pastures, and overgrazing is very severe in these habitats. Thus, the high mountain flora and vegetation are seriously disturbed (personal observation). Since alpine habitats are floristically less well researched, we have insufficient knowledge about the habitats of rare species in the various mountain ranges. We thus strongly recommend more field exploration of high mountain areas to determine the ecology, biology and conservation status of local endemics according to IUCN criteria, and to improve the protection status of the country’s high mountain flora and vegetation.

References

Akhani H 2006. Flora Iranica: facts and figures and a list of publications by K. H. Rechinger on Iran and adjacent areas. - Rostaniha 7/2: 19–61. Al-Shehbaz I & Junussov SJ 2003. Arabidopsis bactriana belongs to Dielsiocharis (Brassicaceae). - Novon 13: 171-172.

Assadi M, Massoumi AA, Khatamsaz M & Mozaffarian V (eds) 1988-2012. Flora of Iran, No. 1-74. - Research Institute of Forest and Rangelands. Tehran. (in Persian).

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Ferrigno JG 1991. Glaciers of the Middle East and Africa – glaciers of Iran. - In: Williams RS & Ferrigno JG (eds.) Satellite image atlas of glaciers of the world. - U.S. Geological Survey Professional Paper 1386-G, United States Government Printing Office, Washington: 31-47. Frey W, Kürschner H & Probst W 1999. Flora and vegetation, including plant species and larger vegetation complexes in Persia. - In: Yarshater E (ed.) Encyclopaedia Iranica 10/1: 43-63. Costa Mesa, California, Mazda Publishers. Gottfried M, Pauli H, Futschik A, Akhalkatsi M, Barančok P, Benito Alonso JL, Coldea G, Dick J, Erschbamer B, Fernández Calzado MR, Kazakis G, Krajči J, Larsson P, Mallaun M, Michelsen O, Moiseev D, Molau U, Merzouki A, Nagy L, Nakhutsrishvili G, Pedersen B, Pelino G, Puscas M, Rossi G, Stanisci A, Theurillat JP, Tomaselli M, Villar L, Vittoz P, Vogiatzakis I & Grabherr G 2012. Continent-wide response of mountain vegetation to climate change. - Nature Climate Change 2/2: 111-115. Grabherr G, Gottfried M & Pauli H 1994. Climate effects on mountain plants. - Nature 369: 448–448. Hedge IC & Wendelbo P 1978. Patterns of distribution and endemism in Iran. - Notes from the Royal Botanic Garden, Edinburgh 36: 441–464. Hedge IC 1968. Dielsiocharis (pp. 320-321), Didymophysa (pp. 87-98) [Cruciferae]. - In: Rechinger KH (ed.) Flora Iranica 57. - Akademische Druck u. Verlagsanstalt, Graz. Hedge IC 1969. Elburzia, a new genus of Cruciferae from Iran. - Notes from the Royal Botanic Garden, Edinburgh 29: 181-184. Jamzad Z 2006. A new species and a new record from Iran. - Iranian Journal of Botany 11/2: 143-148. Kamrani A, Jalili A, Naqinezhad A, Attar F, Maassoumi AA & Shaw SC 2011. Relationships between environmental variables and vegetation across mountain wetland sites, N Iran. - Biologia 66: 76-87. Khalili A 1973. Precipitation patterns of Central Elburz. - Arch Met Geoph Biokl Ser B 21: 215-232. Khassanov F, Noroozi J & Akhani H 2006. Two new species of Allium genus from Iran. - Rostaniha 7(2): 119-130. Klein JC & Lacoste A 1999. Observation sur la végétation des éboulis dans les massifs irano-touraniens: le Galietum aucheri ass. nov. de l’ Alborz central (N-Iran). - Documents Phytosociologiques N. S. 19: 219-228. Klein JC 1982. Les groupements chionophiles de l’Alborz central (Iran). Comparaison avec leurs homolegues d’ Asie centrale. - Phytocoenologia 10: 463-486. Klein JC 1987. Les pelouses xérophiles d’alt itude du franc sud de l’Alborz central (Iran). - Phytocoenologia 15: 253-280. Klein JC 1988. Les groupements grandes ombellifères et a xerophytes orophiles: Essai de synthèse à l’échelle de la région irano-iouranienne. - Phytocoenologia 16: 1-36. Klein JC 1991. Endémisme à l’étage alpin de l’Alborz (Iran). - Flora et Vegetatio Mundi 9: 247-261. Naqinezhad A, Attar F, Jalili A & Mehdigholi K 2010. Plant biodiversity of wetland habitats in dry steppes of Central Alborz. - Australian Journal of Basic and Applied Sciences 4/2: 321-333.

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Naqinezhad A, Jalili A, Attar F, Ghahreman A,Wheeler BD, Hodgson JG, Shaw SC & Maassoumi AA 2009. Floristic characteristics of the wetland sites on dry southern slopes of the Alborz Mts., N. Iran: The role of altitude in floristic composition. - Flora 204: 254–269. Noroozi J & Ajani Y 2013. A new alpine species of Nepeta sect. Capituliflorae (Labiatae) from Northwestern Iran. - Novon DOI: 10.3417/2012022. Noroozi J, Ajani Y & Nordenstam B 2010a. A new annual species of Senecio (Compositae-Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area. - Compositae Newsletter 48: 4–23. Noroozi J, Akhani H & Breckle SW 2008. Biodiversity and phytogeography of alpine flora of Iran. - Biodiversity and Conservation 17: 493-521. Noroozi J, Akhani H & Willner W 2010b. Phytosociological and ecological study of the high alpine vegetation of Tuchal Mountains (Central Alborz, Iran). - Phytocoenologia 40: 293-321. Noroozi J, Pauli H, Grabherr G & Breckle WS 2011. The subnival-nival vascular plant species of Iran: a unique high-mountain flora and its threat from climate warming. - Biodiversity and Conservation 20: 1319-1338. Noroozi J, Willner W, Pauli H & Grabherr G 2013. Phytosociology and ecology of the high alpine to subnival scree vegetations of N and NW Iran (Alborz and Azerbaijan Mts.). - Applied Vegetation Science DOI: 10.1111/avsc.12031. Pauli H, Gottfried M, Dullinger S, Abdaladze O, Akhalkatsi M, Benito Alonso, JL, Coldea G, Dick J, Erschbamer B, Fernández Calzado R, Ghosn D, Holten JI, Kanka R, Kazakis G, Kollár J, Larsson P, Moiseev P, Moiseev D, Molau U, Molero Mesa J, Nagy L, Pelino G, Puşcaş M, Rossi G, Stanisci A, Syverhuset AO, Theurillat J-P, Tomaselli M, Unterluggauer P, Villar L, Vittoz P & Grabherr G 2012. Recent plant diversity changes on Europe’s mountain summits. - Science 336: 353-355. Razyfard H, Zarre S & Fritsch RM 2011. Four new species of Allium (Alliacea) from Iran. - Ann. Bot. Fenn. 48/4: 352-360. Rechinger KH (ed.) 1963-2012. Flora Iranica 1-179. - Akademische Druck- u. Verlagsanstalt und Naturhistorisches Museum Wien. Graz & Wien. Rechinger KH 1968. Zerdana (pp. 307-308) [Cruciferae]. - In: Rechinger KH (ed.). Flora Iranica 57. Akademische Druck u. Verlagsanstalt. Graz. Rechinger KH 1987. Sclerochorton (pp. 366-366) [Umbelliferae]. In: Rechinger, K.H. (ed.) Flora Iranica, No. 162. Akademische Druck u. Verlagsanstalt. Graz. Schweizer G 1972. Klimatisch bedingte geomorphologische und glaziologische Züge der Hochregion vorderasiatischer Gebirge (Iran und Ostanatolien). - In: Troll C (ed.) Landschaftökologie der Hochgebirge Eurasiens. - Franz Steiner Verlag GMBH, Wiesbaden: 221-236. Walther GR, Beißner S, Burga CA 2005. Trends in upward shift of alpine plants. - Journal of Vegetation Science16: 541–548. Zohary M 1973. Geobotanical foundations of the Middle East 2. - Fischer, Stuttgart.

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Chapter 2:

Unknown elements of the subnival-nival flora of Iran

Three papers were published regarding to this subject:

Paper 1: A new annual species of Senecio (Compositae- Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area

Paper 2: Gagea alexii (Liliaceae), a new record from subnival zone of southern Iran with key and notes on sect. Incrustatae

Paper 3: A new alpine species of Nepeta sect. Capituliferae (Labiatae) from Northwestern Iran

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Title of paper: A new annual species of Senecio (Compositae- Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area

Authors: Jalil Noroozi, Yousef Ajani, Bertil Nordenstam

Status: published (2010), Compositae Newsletter 48: 43-62.

Contribution: idea, data collection, data analysis, manuscript writing.

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Title of paper: Gagea alexii (Liliaceae), a new record from subnival zone of southern Iran with key and notes on sect. Incrustatae

Authors: Yousef Ajani, Jalil Noroozi, Igor G. Levichev

Status: published (2010), Pakistan Journal of Botany 42: 67-77

Contribution: data collection, data analysis, manuscript writing.

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Title of paper: A new alpine species of Nepeta sect. Capituliferae (Labiatae) from Northwestern Iran

Authors: Jalil Noroozi, Yousef Ajani

Status: published (2013), Novon 22: 297–303

Contribution: idea, data collection, data analysis, manuscript writing.

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A New Alpine Species of Nepeta sect. Capituliferae (Labiatae) from Northwestern Iran

Jalil Noroozi Department of Conservation Biology, Vegetation and Landscape Ecology, Faculty Centre of Biodiversity, University of Vienna, Rennweg 14, A-1030, Vienna, Austria, and Plant Science Department, University of Tabriz 51666 Tabriz, Iran. Author for correspondence: [email protected]

Yousef Ajani Institute of Medicinal Plants (IMP), Iranian Academic Centre for Education, Culture and Research (ACECR), 55 km Tehran-Qazvin freeway, P.O. Box 31375369, Karaj, Iran. [email protected]

ABSTRACT. A new species, Nepeta sahandica Nor- ing the number of species in this section to eight oozi & Ajani (Labiatae), is described and illustrated (Jamzad, 2009). With the inclusion of the new from the high altitudes of the Sahand Mountains in species here described, the number of species in the northwestern Iran. Morphological differences of this section reaches nine. All species in this section are species from its closest relatives in Nepeta L. sect. either endemics or localized endemics at high Capituliferae (Benth.) Pojark., N. lasiocephala Benth. altitudes of the Zagros and Alborz mountain ranges and N. monocephala Rech.f., are discussed. These (Rechinger, 1982; Jamzad & Assadi, 1984; Jamzad, species differ mainly in their respective habit, basal 1992, 1998, 1999, 2006; Delghandi, 1993; Jamzad and stem leaves, indumentum, and character of bract et al., 2003), but none have extended to the high and calyx. The new species is restricted to the altitudes of the Azerbaijan region in northwestern subnival vegetation zone, associated with unstable Iran, with the exception of the new species scree grounds. Nepeta sahandica is classified as Critically Endangered (CR), based on IUCN Red List described in this paper. criteria. The Sahand refers to a complex of inactive Key words: Iran, IUCN Red List, Labiatae, volcanic mountain peaks in northwestern Iran, east Nepeta, Sahand Mountains. of Lake Urumiyeh (Urmia) and south of Tabriz City, reaching 3707 m.s.m. at its highest summit (Fig. 1). The genus Nepeta L. (Labiatae) is an Irano- The Sahand is surrounded by the Alborz, Zagros, Turanian element and one of the largest genera in Anatolia, and Caucasus mountain ranges and is Iran with ca. 80 species, of which almost 60% are located in the southernmost corner of the Caucasian endemic or subendemic (Rechinger, 1982; Jamzad, region, one of the world’s 25 biodiversity hotspots 2009). It is the second most numerous genus (Myers et al., 2000). Snow remains on the northern represented in the alpine habitats of Iran (21 species) slopes as snowbeds until mid- to late summer, and % with an 86 endemic or subendemic rate (Noroozi et the subnival zone in this region begins ca. 3500 al., 2008). Four Nepeta species have been previously m.s.m. recorded for the subnival-nival flora of Iran (Noroozi The new species was discovered during fieldwork et al., 2011). The high rate of endemism of Nepeta in by the first author in the high altitudes of the Iranian mountains might represent evidence of active Sahand in early August 2010. To identify associ- speciation in the region. In the Flora Iranica,thegenusNepeta was ated species and ecological characters of the 2 divided into 13 sections, with Nepeta sect. Capit- habitat, several phytosociological releve´s (100 m ) uliferae (Benth.) Pojark. characterized by its capi- were sampled from the habitat of the new species, tate, terminal or axillary inflorescences (Rechinger, following the methodology of Braun-Blanquet 1982). Since the publication of Flora Iranica,two (1964). Morphological characters of this species, new species of Nepeta sect. Capituliferae, N. such as the plant habit, root system, and the fresh minuticephala Jamzad and N. natanzensis Jamzad, color of the corolla, were investigated directly in the have been described (Jamzad, 1999, 2006), increas- field. doi: 10.3417/2012022 NOVON 22: 297–303. PUBLISHED ON 24 MAY 2013.

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Figure 1. —A. Topographic map of Iran, indicating the known distribution of Nepeta sahandica Noroozi & Ajani (box), N. monocephala Rech.f. (triangle), and N. lasiocephala Benth. (circle). —B. Topographic close-up of the Sahand Mountains, where the new species N. sahandica is restricted to altitudes above 3500 m.s.m. (darkest shading).

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Nepeta sahandica Noroozi & Ajani, sp. nov. TYPE: Iran. capitate glandular and simple nonglandular tri- East Azerbaijan: Sahand Mtns., 37844 03 N, chomes. Fruit as nutlets 2–2.5 3 0.9 mm, brown, 46829 59 E, 3500–3630 m.s.m., on scree grounds, oblong to elliptic, with 1 prominent median keel. 31 July 2011, J. Noroozi 2598 (holotype, TARI; isotype, W, WU). Figure 2A–F. Distribution and habitat. Nepeta sahandica is known only from the subnival zone of the northern Diagnosis. The new species Nepeta sahandica Noroozi slopes of the Sahand in northwestern Iran, ranging & Ajani differs from N. monocephala Rech.f. and N. from altitudes of 3500–3630 m.s.m. The new species lasiocephala Benth. by the stem leaves restricted mostly to is adapted to unstable scree grounds with a high the lower half of the stem and by the velutinous calyces and percentage of scree cover (ca. 65%) and steep slope leaves. The calyces (6–8 mm) are shorter than those of N. inclination, ca. 308–458. This species thus belongs to lasiocephala (8.5–9 mm), and the lower tooth of the calyx is shorter (2–2.5 mm) than those of both species (ca. 3 and ca. the subnival-nival flora of Iran, a zone with a very 5mminN. monocephala and N. lasiocephala, respectively). high rate of endemism and with narrow geographical In N. sahandica, the bracts are only ciliate and larger (7–8 distributions (Noroozi et al., 2011). A selection of 3 3 1.5–2 mm) versus villous and smaller (3–4 0.5–1 mm) seven phytosociological releve´s from the plant in N. monocephala. community of the new species, following the Perennial, lignified at the base, with multiple, methodology of Braun-Blanquet (1964), is given in ascendent stems that are densely glandular; root Table 1. The vegetation cover of the releve´s was ca. % extended, to 30 cm or more; stems 15–24 cm tall, 15 , and species richness was ca. 10 species per 1.5–2 mm wide at base, not branched or with thin releve´. For more information about the phytosocio- and short branches 4.5–6 cm, all stem parts densely logical aspects of this plant community, see Noroozi covered with capitate glandular trichomes intermixed et al. (2013). with a few simple, nonglandular, multicelluar IUCN Red List category. Vegetation studies in trichomes; stems green shading to violet in upper the high alpine and subnival zones of the Sahand portions of the stem close to the flower heads; stem Mountains show that the new species is restricted to internodes associated with the lowermost leaves 1–3 the highest altitudes, above 3500 m.s.m., and that cm, uppermost internodes 4.5–9.5 cm. Leaves mostly Nepeta sahandica may be very sensitive to global restricted to the lower half of the stems; basal leaves warming as there is no alternative low-temperature petiolate, petioles 9–18 mm; blades 6–13 3 7–13 habitats for upward migration. The Sahand Range is mm, widely ovate to triangular, bases truncate to also under high pressure from summer overgrazing by cordate, margin crenate, apex obtuse; abaxial lamina goats and sheep, whose milk is used to produce the surfaces with prominent veins, covered with capitate region’s famous cheese. In spite of the known high glandular trichomes intermixed with sparse to rate of local endemism in these mountains, there are moderate simple, nonglandular, multicellular tri- no appropriate control and protection plans. There- chomes, velutinous, adaxial lamina surfaces with fore, N. sahandica is assessed as Critically Endan- few capitate, glandular trichomes, but with dense, gered (CR), according to IUCN Red List criteria simple, nonglandular, multicellular trichomes; stem (IUCN, 2001). leaves subsessile, blades 6–14 3 4–8 mm, ovate to oblong, bases cuneate, margin denticulate, apex Phenology. The new species has been seen in acute. Inflorescences capitate and terminal, 14–18 flower from mid-July to mid-August. 3 14–18 mm; pedicels ca. 1 mm long; bracts linear, 7–8 3 1.5–2 mm, violet in middle portion, 1 vein Etymology. The epithet of the new species refers evident dorsally, with sparse, simple, nonglandular, to its type locality in the Sahand Mountains. multicellular trichomes, velutinous, margin ciliate. Flowers with the calyx 6–8 3 1 mm, tubular, throat Taxonomic affinities. The presumed closest rel- oblique, with capitate glandular and dense, simple, atives of the new species include Nepeta monoce- nonglandular trichomes, velutinous thoughout, violet phala and N. lasiocephala, and both are recorded in at the least upper and middle portions; calyx teeth from the Zagros mountain ranges, extending to the lanceolate, acuminate or acute, with the lower tooth south and southeast of the Sahand (Fig. 1). Nepeta 2–2.5 mm long, the upper tooth 1.5–2 mm long; monocephala is known only from the type collection corolla tubular, blue-violet, 12–13 mm, the corolla in northern Zagros, collected in 1968 from the Chang tube exerted from the calyx, the outer surface covered Almas Mountain at altitudes between 2200 and 2500 by short, simple, nonglandular trichomes, dorsal m.s.m. (Rechinger, 1982); it has not been recorded or portions of the upper lip densely covered with collected since then. The description of N. mono-

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Figure 2. A–F. Nepeta sahandica Noroozi & Ajani. —A. Inflorescence. —B, C. Entire plant. —D. Calyx. —E. Corolla. —F. Nutlet. G–J. Nepeta lasiocephala Benth. —G. Inflorescence. —H, J. Entire plant. Photos A–C taken by J. Noroozi in the Sahand, 3550 m.s.m.; photos D–F taken by Y. Ajani from the TARI holotype, J. Noroozi 2598 (TARI); photos G–J taken by J. Noroozi and Y. Ajani in the Hezar Mountains, 4450 m.s.m.

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Table 1. Species composition of the habitat for Nepeta sahandica, based on seven phytosociological releve´s from the subnival zone of the Sahand, at the northern slope of its highest peak, on Kamal Daghi (3707 m.s.m.), and for N. lasiocephala, based on two releve´s from the subnival zone of the Hezar Mountains, on the northern slope of its highest peak (4465 m.s.m.). The symbols show the cover-abundance of each species, according to the method of Braun-Blanquet (1964), where 2 ¼ 5%–25%;1¼ 1%–5%; þ¼, 1%; and r ¼ , 5cm2. Each releve´ area is 100 m2.

Sahand Hezar Releve´ number 1088 1089 1090 1091 1092 1093 1095 938 939 Altitude (m.s.m.) 3538 3550 3553 3567 3571 3592 3622 4455 4443 Slope aspect NNNNNNNENENE Slope inclination (degrees) 40 45 40 45 45 35 30 20 20 Plant cover (%) 201515101015305 10 Scree coverage (%) 454040858060759285 Solid rock (%) 303540001502 3 Bare ground (%) 5 10 5 5 10 10 5 1 2 Species richness 16 9 11 8 9 8 13 7 11

Nepeta sahandica 1112111 Didymophysa aucheri Boiss. 1111111 Bromus tomentosus Trin. 1111122 Ziziphora clinopodioides Lam. 1111111 Senecio taraxacifolius (M. Bieb.) DC. 1 1 1 þ 1 — 1 Poa araratica Trautv. 1 1 1 1 — þ þ Festuca alaica Drobow 1 — 1 þ 1 1 1 Dracocephalum aucheri Boiss. 1 2 1 — þ 1 — Arabis caucasica Willd. þ — þ — — þ 1 Galium hyrcanicum C. A. Mey. — — þ þ 1 — 2 Saxifraga sibirica L. þ þ — — 1 — — Pedicularis caucasica M. Bieb. þ — — — — — þ Erysimum gelidum Bunge þ — — — — — 1 Erigeron caucasicus Steven þ — — — — — 1 Helichrysum psychrophilum Boiss. — — — — — — þ Crepis frigida (Boiss. & Balansa) Babc. þ — þ — — — — Nepeta menthoides Boiss. & Buhse 1 — — — — — — Thymus kotschyanus Boiss. & Hohen. ex Boiss. þ — — — — — — Artemisia L., sp. indet. — r — — — — —

Nepeta lasiocephala 11 Astragalus melanodon Boiss. 11 Artemisia persica Boss. þ 1 Asperula glomerata (M. Bieb.) Griseb. subsp. þ 1 condensata (Ehrend.) Ehrend. Potentilla nuda Boiss. 1 þ Scrophularia subaphylla Boiss. þ þ Gagea alexii Ali & Levichev þ — Ranunculus eriorrhizus Boiss. & Buhse — 1 Senecio subnivalis Y. Ajani, J. Noroozi & B. Nord. — þ Veronica kurdica Benth. subsp. filicaulis (Freyn) — þ M. A. Fisch. Psychrogeton alexeenkoi Krasch. — þ Silene daenensis Melzh. — þ

cephala is thus based only on the type specimens in phological characters. The basal leaves of these the W and IRAN herbaria. We have seen live plants species are identical in shape, margin, apex, and of N. lasiocephala (Fig. 2G–J) on the highest summit basal parts. The major morphological differences of the Hezar Mountains (4465 m.s.m.) in southern between these species include the plant habit, stem Iran, and we have also examined the type specimens trichome types and indumentum, leaf sizes and of both N. lasiocephala and N. monocephala in the W indumentum, bract sizes, and calyx sizes and herbarium. These three species share several mor- indumentum, as well as the corolla shape and length

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Table 2. Morphological differences among Nepeta sahandica, N. monocephala, and N. lasiocephala.

Characters N. sahandica N. monocephala1 N. lasiocephala2 Habit ascendant straight ascendant or deflexed Stem trichome types; dense capitate glandular subsessile glandular trichomes dense capitate glandular indumentum trichomes intermixed with a intermixed with simple trichomes intermixed with few simple nonglandular nonglandular multicellular simple nonglandular multicellular ones; densely ones; basally villous, multicellular ones; villous glandular glabrescent above Basal leaf sizes (mm) 6–13 3 7–13 15–20 3 15–20 5–10 3 5–10 Stem leaf sizes (mm), 6–14 3 4–8, mostly in lower 2–4 3 0.5–1, stems nearly 5–10 3 5–10, leaves equally position, shape, margin half, but some also in upper naked (subnude), leaves distributed, ovate, crenate half of the stem, leaves basal, lanceolate, ovate to oblong, denticulate denticulate, or entire Leaf indumentum velutinous, denser adaxially densely villous, more so densely lanate, more so than abaxially abaxially than adaxially abaxially than adaxially Bract size (mm), 7–8 3 1.5–2, glabrate, ciliate 3–4 3 0.5–1, villous 8 3 2.5–3, villous in upper indumentum at margin portions Calyx indumentum velutinous villous villous in upper portions Calyx size (mm) 6–8 3 1 7–8 3 1.5–2 8.5–9 3 1 Calyx lower tooth size 2–2.5 3 5 (mm) Calyx upper tooth size 1.5–2 1.5–2 2.5–3 (mm) Corolla shape tubular tubular funneliform Corolla length (mm) 12–13 12–13 11 Nutlet shape oblong to elliptic ovate to elliptic oblong Nutlet size (mm) 2–2.5 3 0.9 1.3 3 0.8 1.9 3 0.9 1 Data according to Rechinger (1982) and isotype Iranshahr & Dezfoulin E13207 (W). 2 Data according to Rechinger (1982), isotype Kotschy 757 (W), and field studies.

(see Table 2). Nutlet shape and size are also is no information available for the habitat of N. significant characters for the taxonomy of the tribe monocephala because this species is known only from Nepeteae (Budantsev, 1993; Budantsev & Lobova, the type collection and has not been recorded since, 1997). Distributional and ecological characters of the despite some general, although not detailed, botan- three similar species are compared in Table 3. Nepeta ical surveys by local botanists (H. Maroofi, pers. natanzensis is another member of Nepeta sect. comm.). The type locale for N. monocephala on the Capituliferae, but it differs greatly from the discussed Chang Almas Mountain has not been botanically well species in its clearly lignified caudex and basal parts of the stem and its spine-shaped calyx. explored, and its precise location is not known. It is The habitat of Nepeta lasiocephala was studied in likely that this species may be found again with the Hezar Mountains, in southern Iran, with sampling botanical investigation. There is no doubt that this of two phytosociological releve´s. Table 1 compares species is very rare and locally endemic, and further the ecology and species composition of this species botanical exploration of its distribution, habitat, and with N. sahandica, showing the data of releve´s. There ecology is highly recommended.

Table 3. Distributional and ecological characters associated with Nepeta sahandica, N. monocephala, and N. lasiocephala.

Characters N. sahandica N. monocephala N. lasiocephala Geographical distribution Sahand Mtns. northern Zagros (Chang Almas central, south, and Mtn.) southeastern Zagros Altitudinal distribution 3500–3630 2200–2500 2000–4465 (m.s.m.) Habitat scree grounds of subnival zone no data scree grounds of subalpine to subnival zones Threat status (IUCN Red Critically Endangered (CR) Data Deficient (DD) (its Near Threatened (NT) List criteria) presence has not been recorded for 44 years)

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Acknowledgments. We are grateful to Jose´ Luis Commission. IUCN, Gland, Switzerland, and Cambridge, Ferna´ndez-Alonso, Sara Fuentes Soriano, and Victo- United Kingdom. Jamzad, Z. 1992. Two new species from Labiatae in Iran. ria Hollowell for their constructive comments and Iranian J. Bot. 5(2): 69–74. edits on a former version of the manuscript, and to Jamzad, Z. 1998. A new species of Nepeta L. (Labiatae) Johannes Lundberg for editing the English text. Ernst from Iran. Iranian J. Bot. 7(2): 249–253. Vitek, curator of the W herbarium, is acknowledged Jamzad, Z. 1999. Two new species of Nepeta L. (Labiatae) for his assistance in investigating the type specimens from Iran. Iranian J. Bot. 8(1): 43–48. Jamzad, Z. 2006. A new species and a new record from Iran. of the related species, and Bruno Wallno¨fer is Iranian J. Bot. 11(2): 143–148. acknowledged for his help in determining the Jamzad, Z. 2009. Notes on the genus Nepeta L. (Lamiaceae- trichome types of these species. Traveling and Nepetoideae). Iranian J. Bot. 15(2): 141–145. subsistence costs for this study were covered by Jamzad, Z. & M. Assadi. 1984. New species of the genera Nepeta and Ajuga (Labiatae) from Iran. Iranian J. Bot. funds from the Global Observation Research Initia- 2(2): 95–102. tive in Alpine Environments (GLORIA) coordination Jamzad, Z., M. Ingrouille & M. S. J. Simmonds. 2003. Three (University of Vienna and Austrian Academy of new species of Nepeta (Lamiaceae) from Iran. Taxon 52: Sciences/Institute of Mountain Research), the asso- 93–98. ciation GLORIA-International, Vienna, and the Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. Fonseca & J. Kent. 2000. Biodiversity hotspots for Austrian Federal Ministry of Science and Research. conservation priorities. Nature 403: 853–858. Noroozi, J., H. Akhani & S. W. Breckle. 2008. Biodiversity Literature Cited and phytogeography of the alpine flora of Iran. Biodivers. & Conservation 17(2): 493–521. Braun-Blanquet, J. 1964. Pflanzensoziologie: Grundzu¨ge Noroozi, J., H. Pauli, G. Grabherr & S. W. Breckle. 2011. der Vegetationskunde. Springer-Verlag, Vienna. The subnival-nival vascular plant species of Iran: A Budantsev, A. L. 1993. Ultrastructural features of fruit unique high-mountain flora and its threat from climate surface in some genera of the tribe Nepeteae (Lam- warming. Biodivers. & Conservation 20(6): 1319–1338. iaceae). Bot. Zhurn. 78: 100–108. Noroozi, J., W. Willner, H. Pauli & G. Grabherr. 2013. Budantsev, A. L. & T. A. Lobova. 1997. Fruit morphology, Phytosociology and ecology study of the high-alpine to anatomy and taxonomy of tribe Nepeteae (Labiatae). subnival scree vegetation of N and NW Iran (Alborz and Edinburgh J. Bot. 54(2): 217–229. Azerbaijan Mts.). Appl. Veg. Sci. DOI: 10.111/avsc. Delghandi, M. 1993. Nepeta leucostegia (Labiatae), a new 12031. record for the flora of Iran. Iranian J. Bot. 6(1): 149–151. Rechinger, K. H. 1982. Nepeta. Pp. 108–216 in K. H. IUCN. 2001. IUCN Red List Categories and Criteria, Rechinger (editor), Flora Iranica, Vol. 150. Akademische Version 3.1. Prepared by the IUCN Species Survival Druck-u. Verlagsanstalt, Graz.

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Chapter 3:

Vegetation of the alpine and subnival-nival zones of N and NW Iran

One paper was published regarding to this subject:

Paper: Phytosociology and ecology of the high alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.)

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Title of paper: Phytosociology and ecology of the high alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.)

Authors: Jalil Noroozi, Wolfgang Willner, Harald Pauli & Georg Grabherr

Status: published (2013), Applied Vegetation Science DOI: 10.1111/avsc.12031.

Contribution: idea, data collection, data analysis, manuscript writing.

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Applied Vegetation Science && (2013) Phytosociology and ecology of the high-alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.) Jalil Noroozi, Wolfgang Willner, Harald Pauli & Georg Grabherr

Keywords Abstract High mountains; Plant community; Southwest Asia; Syntaxonomy; Vascular plants Questions: The vegetation of high-alpine and subnival scree habitats in Iranian mountains has been poorly investigated so far despite the large Nomenclature variety of narrowly distributed vascular plant species and the expected Rechinger (1963–2012) vulnerability of these ecosystems to global warming. Which plant

Received 08 May 2011 communities occupy these ecosystems and what is their syntaxonomic Accepted 13 January 2013 position? Which environmental factors determine the species composition of Co-ordinating Editor: Joop Schaminee these habitats? Location: Alborz and Azerbaijan Mountains in N and NW Iran. Noroozi, J. (corresponding author, noroozi. Methods: A total of 141 phytosociological releves were collected from 3200 to [email protected]) & Grabherr, G. ([email protected]): Department of 4800 m a.s.l. This data set was classified using TWINSPAN, and the numerical Conservation Biology, Vegetation and classification was translated into a syntaxonomic system. Floristic differences Landscape Ecology, University of Vienna, between vegetation types were evaluated with detrended correspondence anal- Rennweg 14, 1030, Vienna, Austria ysis (DCA). We determined the means and SD of measured environmental and Willner, W. ([email protected]): vegetation parameters for all associations. Differences in the major environmen- Vienna Institute for Nature Conservation & tal parameters among associations and alliances were analysed using ANOVA Analyses, Giessergasse 6/7, 1090, Vienna, and post-hoc tests. Moreover, we determined the mean cover percentage of life Austria Pauli, H. ([email protected]): Institute forms in all associations. for Interdisciplinary Mountain Research (IGF) of Results: All high-alpine and subnival scree communities are arranged in one the Austrian Academy of Sciences, c/o class (Didymophyso aucheri-Dracocephaletea aucheri), two orders (Physoptychio University of Vienna, Rennweg 14, 1030, Vienna, Austria gnaphalodis-Brometalia tomentosi, Didymophysetalia aucheri), three alliances (Elymo longearistati-Astragalion macrosemii, Erigerontion venusti, Didymophysion aucheri) and ten associations, which are new to science, except for one association. The territory of the class extends from Alborz to NW Iran and probably to E Anatolia, Transcaucasia and the Zagros Mountains. Altitude, aspect and edaphic qualities are the major ecological factors influencing the species composition and vegetation mosaic. Conclusions: Our study introduces a formal syntaxonomic classification of the scree vegetation at high altitudes in Iran, thus providing a scheme for ongoing ecological surveys and monitoring programmes to assess the impacts of climate warming and of human land use on these unique ecosystems.

present in the alpine belt of the Iranian mountains, more Introduction than 50% are endemic or sub-endemic to the country In spite of the vast and diverse mountain areas of Iran, the (Noroozi et al. 2008). A total of 151 species reach the high-alpine and subnival–nival areas have a highly subnival–nival belt, and of the 51 species with mainly scattered distribution throughout the country, and the subnival–nival distribution, 68% are endemic to Iran, most rate of endemic vascular plants with narrow geographic of them being found in only one or a few mountain ranges distribution is very high in comparison to lower altitudes (Noroozi et al. 2011). The alpine ﬂora of Iran has (Noroozi et al. 2011). Of the ca. 700 vascular plant species Irano-Turanian origin (Zohary 1973; Klein 1982, 1991;

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Scree vegetation of Iran J. Noroozi et al.

Frey et al. 1999; Noroozi et al. 2008, 2011). It exhibits the Study area strongest floristic relationships with Anatolia, the Hindu Kush and the Caucasus, which have 23%, 20% and 19% Alborz is a W–E ranging mountain chain, located in N Iran, of species in common with the high mountains of Iran, along the southern shore of the Caspian Sea, with Mt. Da- respectively (Noroozi et al. 2008). mavand, at 5671 m a.s.l., being the highest peak in the A consistent large-scale classification of floristically country. In NW Iran there are two isolated high volcanic defined vegetation types is an important tool for ecologi- mountains: Sabalan, which is the third highest peak in cal research, bioindication, vegetation monitoring, Iran, at 4811 m a.s.l. and having an active glacier near its conservation strategies and legislation (Dengler et al. peak, and Sahand, a complex of several volcanic summits 2008). The first vegetation survey on high elevations of with a maximum elevation of 3707 m a.s.l. (Fig. 1). These Central Alborz dates back to Kotschy (1861a,b), Buhse two mountains in the NW part of Iran are located between (1899a,b) and Gilli (1939, 1941). An overview of the four main mountain ranges: Alborz in the east, Zagros in flora and phytogeography of the alpine and subnival– the south, the Caucasus in the north and Anatolia in the nival zone in the Iranian mountains was provided by west. Noroozi et al. (2008, 2011). Phytosociological studies of All study sites are located on volcanic and volcaniclastic the alpine zone of Iran were carried out only in Central formations with acidic soils. The main body of the Damav- Alborz, and the syntaxonomic inventory of Iran is still far and volcano, remarkably around the summit including the from complete. A first syntaxonomic treatment of the sampling area, is composed of trachyandesitic lava flows sub-alpine and alpine vegetation of Central Alborz was and pyroclastic sediments of Quaternary age (Davidson published by Klein (1982, 1987, 1988), who introduced et al. 2004). Tuchal summit and its S and N flanks are con- three classes, which are, however, not validly published: structed of Eocene acidic volcanic tuffs and tuffaceous silt- ‘Prangetea ulopterae’ (sub-alpine heaths and tall herb vege- stones/shales (Jamshidi & Afsharian Zadeh 1993). Sabalan tation), ‘Onobrychidetea cornutae’ (lower alpine thorn cush- is a Plio-Quaternary volcano, mainly composed of potas- ion vegetation) and ‘Oxytropidetea persicae’(higheralpine sium-rich calc-alkaline andesitic rocks (Innocenti et al. dwarf shrub vegetation). Noroozi et al. (2010) presented 1982). Sahand volcano is a volcanic complex that has a new classification of the snowbed and thorn cushion formed through two major episodes of volcanic activity: grasslands of the alpine zone of Central Alborz and sug- during Middle-Upper Miocene and Plio-Quaternary. The gested that the above-mentioned classes should be recon- studied plots are localized over the Plio-Quaternary rocks sidered. A synopsis of plant communities and their mostly comprised of calc-alkaline dacitic and andesitic elevational patterns on the southern slopes of Central rocks (Innocenti et al. 1982; Behrouzi et al. 1997). The soil Aborz from the low-montane to the high-alpine zone, is of all these volcanic areas is constituted of lithosols (igne- presented in Akhani et al. (2013). The only phytosocio- ous rocks) (Dewan et al. 1961). logical work on scree vegetation of subnival areas of Iran There are glaciers in the higher elevations of the three was published by Klein & Lacoste (2001), describing one highest mountains of Iran, i.e. Damavand, Alamkuh and association (Galietum aucheri) based on 14 phytosociologi- Sabalan (Ferrigno 1991; Fig. 1). The present snowline in cal releves from Central Alborz. The high mountains of Alborz and NW Iranian mountains lies between 4000 and NW Iran are even less known, and our study presents 4200 m a.s.l. (Schweizer 1972), and the upper limit of vas- the first vegetation data for this area. cular plants in Central Alborz and Sabalan is 4800 and We describe the alpine and subnival–nival scree 4500 m a.s.l., respectively (Noroozi et al. 2011). It is vegetation of the Central Alborz and the mountains of assumed that during the Pleistocene the climatic snowline Azerbaijan (NW Iran), based on 141 of our own releves in the Iranian mountains was 1200–1800 m lower than at plus the afore-mentioned 14 releves of Klein & Lacoste present (Wright 1962). Therefore, the upper limit of vascu- (2001). The purpose of this research was to address the lar plants might have been around 1500 m below its pres- following questions: (1) which major plant community ent level. types can be distinguished in high-altitude screes in Cen- The higher altitudes of Alborz are affected by the tral Alborz and the mountains of NW Iran, and how can north-westerly flow of polar air masses, and annual pre- their compositional variation be adequately represented cipitation reaches almost 1000 mm (Khalili 1973). The in a syntaxonomic classification; (2) what are the ecologi- summer is arid, hot and sunny, with intense radiation cal, life-form and biogeographical characteristics of these most of the time. Both annual and diurnal amplitudes of syntaxa; and (3) what is the syntaxonomic relationship temperature can be very high, in particular near the soil between these vegetation types and similar habitats of surface. Although the duration of snow cover might be Anatolia, the Caucasus, the Balkan Peninsula and the an important factor determining vegetation patterns, European Alps? there are no data available to discuss this factor for the

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Fig. 1. The study areas in N and NW Iran and the number of releves recorded by the first author in each mountain area. Damavand (5671 m a.s.l., highest peak of Iran), Alamkuh (4850 m a.s.l., second highest peak of Iran), Sabalan (4811 m a.s.l., third highest peak of Iran), Tuchal (3966 m a.s.l.), Sahand (3707 m a.s.l.). The 14 releves of Klein & Lacoste (2001) are from Central Alborz [Alamkuh, Azadkuh and Damavand, with 11, 2 and 1 releves, respectively]. communities of this study. Based on the Global Biocli- vegetation cover, high percentage of scree on the ground matic Classification System (GBC) developed by Rivas- and distinct species composition. It was decided at the out- Martınez et al. (1997, 1999), the high plateaus of NW set of the field sampling to record scree vegetation with Iran and most of Alborz (except the Hyrcanian region) plot sizes of 10 m 9 10 m, thorn cushion grasslands at belong to the mediterranean macrobioclimate, with long 5m9 5 m, and snowbeds at 2 m 9 2 m. However, after summer droughts, but colder winters compared to the a preliminary analysis of the data, we found that some of mediterranean region (Djamali et al. 2011). Sahand has the communities on scree ground rather belong to the driest climate among the studied mountains (medi- snowbed vegetation in terms of species composition, while terranean xeric-continental), Central Alborz has a more some stands initially classified as thorn cushion grassland humid climate (mediterranean pluviseasonal continen- have a closer floristic relationship to scree vegetation tal), and Sabalan has the least continental bioclimate (data not shown). Therefore, the releves selected for the (mediterranean pluviseasonal-oceanic). present study vary in plot size between 25 and 100 m². This is, however, still within the extremes of the plots of Klein & Lacoste (2001), which vary from 5 to 400 m².Our Methods target vegetation types do not extend to lower altitudes From 2003 to 2011, a total of 141 releves were collected in because the scree grounds of the sub-alpine zone are the scree vegetation of the Central Alborz and NW Iran, almost fully occupied by communities of the class Prangetea focusing on altitudes between 3200 and 4800 m a.s.l., and ulopterae Klein 1987 nom. inval., i.e. tall herb vegetation following the traditional sampling strategy in the Braun– dominated by umbelliferous plants, which have a totally Blanquet approach (Dengler et al. 2008). The releves were different physiognomy and floristic composition (Klein taken in open plant communities growing on unstable or 1987, 1988). For each plot, elevation, geographic stable stony and scree substrates. Usually, these vegetation coordinates, aspect, slope inclination, as well as percentage types are easily distinguished from alpine snowbeds and of open scree (gravel or larger stones), solid rock, bare thorn cushion grasslands (Noroozi et al. 2010) by their low ground (open soil) and vegetation cover were recorded.

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Species cover was recorded using a modified Braun–Blan- lower but a character species at the higher level, or (2) if quet scale (r: <5cm², +: <1%, 1: 1–5%, 2: 5–25%, 3: 25– it was diagnostic at a low and a high level but not at the 50%, 4: 50–75%, 5: 75–100%). Due to the low relevance intermediate level (e.g. at the association and order level of bryophytes and lichens in the studied vegetation types but not at the alliance level), or (3) if it was considered (see Photo S1–S17), which is probably caused by the dry- as a transgressive character species. For the last, various ness of the growing season, only vascular plants were definitions exist (e.g. Dengler 2003: 97; Luther-Mosebach recorded. et al. 2012: 406). We considered a species as a transgres- For data storage and table work, the programs TURBO- sive character species of a syntaxon if its fidelity for this VEG and JUICE were used. The releves were first classified syntaxon was lower than for the next higher level. Note using the TWINSPAN algorithm, using percentage cover that following the proposal of Willner et al. (2009), total values of 0, 5, 25 and 50 as cut-off levels. Associations were cover values <0.1% were set to 0.1 for the calculation of delimited following the proposals of Willner (2006). The TCR. final delimitation included the re-arrangement of a single We determined the means and SD of measured environ- releve according to the summarized cover of the diagnostic mental and vegetation parameters (altitude, aspect, incli- species (see Willner 2011). Floristic differences between nation, species richness per plot, Shannon diversity index vegetation types were also evaluated with detrended and Smith–Wilson evenness index (Smith & Wilson 1996), correspondence analysis (DCA) using Canoco for Windows vegetation cover, open scree cover, bare ground cover, v. 4.5. Associations were arranged into higher units (alli- solid rock cover) for all associations. Differences in the ances, orders) by taking into account the TWINSPAN and major environmental parameters among associations and DCA result, but the TWINSPAN hierarchy was not strictly alliances were analysed using ANOVA and Games–Howell followed. Instead, we aimed at achieving a classification post-hoc tests and illustrated with box plots. We also deter- that maximizes the number and fidelity of absolute charac- mined the mean cover percentage of life forms in all associ- ter species for the higher units, although a strict statistical ations using the average cover percentage corresponding evaluation of this criterion is hardly possible with the avail- to each value of the modified Braun–Blanquet scale. We able data (see Willner et al. 2009; Luther-Mosebach et al. assigned all taxa to six categories of life forms: therophyte, 2012). The distinction between differential and character small hemicryptophyte (ca. 30 cm), graminoid, cushion chamaephyte and non- available information about the habitat and geographical cushion chamaephyte. distribution of the species. A species was considered as Flora Iranica (Rechinger 1963–2012) was used as the diagnostic for a syntaxon if its constancy ratio (CR) was main source for identification and nomenclature of plants. 2, i.e. if the constancy of the species in a unit was at least Furthermore, several other monographs were used to twice as high as in any other unit of the same rank within identify some taxa such as Elymus longearistatus (Davis the next higher hierarchical level (Dengler 2003). A spe- 1985), Erysimum (Polatschek 2011) and Festuca alaica (Al- cies with CR < 2 was considered diagnostic if its total cover exeev 1979). Information on the phytogeography of diag- (=average cover taking also into account zero values) was nostic species was obtained from various literature sources. >15% and its total cover ratio (TCR) was 2, i.e. if its total Specimens are preserved at the personal herbarium of the cover in this unit was at least twice as high as in any other first author. For nomenclature of syntaxa, the Interna- unit of the same rank within the next higher hierarchical tional Code of Phytosociological Nomenclature (Weber level. Species cover in individual releves was approxi- et al. 2000) was applied. mated by taking the average cover corresponding to each value of the Braun–Blanquet scale. We preferred the TCR Results and discussion over other fidelity measures, such as the Phi coefficient General floristic and physiognomic features of association, because it includes information on species cover, and provides a simple criterion for distinguishing The number of species recorded in all plots totalled 101, non-diagnostic, diagnostic and strongly diagnostic species where ca. 70% of the species belong to six families (Astera- (Willner et al. 2009). However, because of the unavoid- ceae: 19 species, Poaceae: 13, Brassicaceae: 11, Labiatae: able inaccuracy of the calculated total cover values, we ten, Scrophulariaceae: eight, Fabaceae: seven species). used a combination of CR and TCR for the definition of Constancy above 1% (i.e. at least two occurrences) was diagnostic species. Species with a total cover <1% were reached for 77 species. The highest constancies within the only exceptionally considered as diagnostic. If a species whole data set were (values in brackets): Bromus tomentosus reached the threshold for a diagnostic species at several (65%), Poa araratica (37%), Dracocephalum aucheri (35%), hierarchical levels, it was only considered at the lowest Festuca alaica (35%), Ziziphora clinopodioides (32%) and Did- level, except (1) if it was a differential species at the ymophysa aucheri (28%).

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Almost 80% of the plant species in our data set are Syntaxonomic assignment to a class hemicryptophytes, followed by chamaephytes (14%) and therophytes (6%). In comparison to the alpine thorn cush- Klein & Lacoste (2001) classified the subnival–nival ion grasslands of the Central Alborz (Noroozi et al. 2010), scree vegetation of Central Alborz as provisional class the percentage of hemicryptophytes is higher while the ‘Didymophysetea aucheri’. A synoptic table of the most percentage of chamaephytes and therophytes is lower, and important alliances of scree vegetation in the European there are no geophytes in our plots. Thus, our result is sim- Alps, the Balkan Peninsula, the Caucasus, Anatolia, as ilar to previous observations that the percentage of hemi- well as the alliances described in the present paper, sup- cryptophytes increases from lowlands to high elevations in ports this proposal (Table 2, Appendix S1): the only Iranian mountains (Noroozi et al. 2008, 2010; Naqinezhad scree species present in all five regions is Oxyria digyna, et al. 2009), as also found in the Hindu Kush (Agakhan- which occurs in two associations of our study area janz & Breckle 1995) and in Mediterranean mountains (Table 1). Other species with a wide geographic distribu- (Kazakis et al. 2007; Fernandez Calzado & Molero Mesa tion are Arabis alpina agg. (including A. caucasica), which 2011). Hemicryptophytes can be classified into three is recorded in all regions except the Caucasus (sic!), groups: rosettes/small hemicryptophytes (54%), tall hemi- Cerastium cerastoides, which is rather a snowbed species cryptophytes (12%) and graminoids (14%). and not present in our releves, and Draba siliquosa. Sedum tenellum is a frequent species of scree vegetation in the Caucasus (Onipchenko 2002) but rarely occurs in Numerical classification and DCA ordination Anatolia and Iran. Seven more species are common to The alliances described below correspond with the first Iran and the Caucasus (Alopecurus dasyanthus, Carum cau- and second level of division in the TWINSPAN classifica- casicum, Lamium tomentosum, Minuartia recurva, Saxifraga tion (Table 1). Cluster 1-1 comprises the communities of sibirica, Scrophularia variegata, Senecio taraxacifolius), and the upper limit of vascular plants in Central Alborz, with two species are common to Iran and Anatolia (Jurinella low vegetation cover and low species richness. They are moschus, Pisum formosum). In contrast, 32 species are classified as alliance Didymophysion aucheri. The releves of unique to Iranian scree vegetation. Of 55 character spe- Klein & Lacoste (2001) are all accommodated within this cies of the Thlaspietea rotundifolii or subordinate syntaxa cluster. Cluster 1-0 comprises scree vegetation in the high- (according to Valachovic et al. 1997) present in the Alps alpine belt of Central Alborz and communities interposing and the Balkan, only four species occur in the Caucasus, alpine thorn cushion grasslands with subnival scree habi- three reach Anatolia, and two the Iranian mountains. A tats in the same region. This cluster represents the alliance single class comprising the scree vegetation of Europe as Elymo longearistati-Astragalion macrosemii. Cluster 0 includes well as of W Asia seems rather ill defined. There is some the scree communities of the Sahand and Sabalan Mts. It is floristic relationship between the Caucasus and NW classified as alliance Erigerontion venusti. Although this alli- Iran, but uniting these two regions into one class would ance was separated from the others at the first TWINSPAN neglect the long list of common differential species level, the communities of NW Iran have a considerable flo- between the Caucasus and the European mountains ristic, physiognomical and ecological similarity to the com- (see Appendix S1). The scree vegetation of Iranian munities of the Elymo-Astragalion (cluster 1-0), including mountains can certainly not be accommodated within high cover and constancy of Bromus tomentosus, Ziziphora the Heldreichietea Quezel ex Parolly 1995 of Anatolia (Pa- clinopodioides and Poa araratica. Therefore, we propose to rolly 1995, 1998), and therefore the description of a join these two alliances into one order (see below). The new class seems unavoidable. The distribution patterns associations described in this study are separated at the of its character species (which are described in the next third or fourth level of the TWINSPAN classification sections) suggest that the territory of this class covers (Table 1). the whole Atropatenian subprovince (sensu Takhtajan In the DCA diagram (Fig. 2), the communities of NW 1986), which includes the arid and semi-arid parts of Iran (Erigerontion venusti) are clustered on the left and the Transcaucasia, E and SE Turkey, S Armenian highlands, communities of the high-alpine and lower subnival zone NW Iran and the Alborz mountains. This subprovince is of Central Alborz (Elymo-Astragalion) on the right. Within one of the most active centres of speciation in W Asia both alliances, the communities with high cover of grasses (Takhtajan 1986). Thus, the concept of the class as out- are concentrated in the centre of the diagram. The commu- lined by Klein & Lacoste (2001) can be confirmed. The nities of the higher subnival–nival zone of Central Alborz class may also extend to the Zagros Mountains, but (Didymophysion) are clustered in the upper part of the dia- more data from this region are required to confirm this gram. assumption.

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Table 1. Synoptic table of the scree communities in N and NW Iran. Values are percentage constancies. The constancy values of diagnostic species are shaded, character species are additionally framed. A dashed frame indicates that the character species are only present in certain sub-associations. In the case of transgressive character species, the higher level is indicated with light shading. If a species has a total cover of >15%, the constancy value is in bold. Companion species that occur in less than ten plots are not shown. Association numbers are the same as in Tables 3 and 4, and Fig. 2. For the association names, see Table 4.

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Table 1. (Continued)

Description of vegetation units of the Atropatenian subprovince (sensu Takhtajan 1986) and probably also of Zagros. The communities of this class The plant communities are arranged in two orders, three are distinguished from alpine snowbed and thorn cushion alliances and ten associations. From these associations, one grasslands (Noroozi et al. 2010) by low vegetation cover, was previously described in Klein & Lacoste (2001), eight high proportion of open scree cover and different species are formally described as new, and one is only presented as composition. The character species of the class are Didymo- a provisional association. The mean and SD of the mea- physa aucheri and Dracocephalum aucheri. Both species typi- sured ecological factors and vegetation parameters for all cally dwell on scree ﬁelds and are indicator species of the associations and alliances, as well as the result of the ANO- subnival–nival zone. Didymophysa aucheri (Photo S1) is dis- VA and post-hoc tests, are given in Appendix S2, and the tributed in E Anatolia, Transcaucasia, NW Iran, Zagros and differences among the units are illustrated in Fig. 3. Mean Alborz (Hedge 1968), and Dracocephalum aucheri (Photo proportional cover of life forms in the ten associations is S1) from SE Anatolia to NW Iran and Alborz (Rechinger shown in Fig. 4. The character and differential species are 1982b). The distribution of these two species (especially listed in alphabetical order in the text by decreasing con- Didymophysa aucheri) probably also determines the geo- stancy in Table 1. A synoptic table with the total cover and graphical borders of the class. Alopecurus textilis is equally ﬁdelity degree of all species is given in Appendix S3. For common in alpine thorn cushion grasslands of the Acantho- type releves, see Table 3. limion demawendici (Noroozi et al. 2010) and occurs from Lebanon, Anatolia, Iraq, Iran to Turkmenistan (Bor 1970). 1- Didymophyso aucheri-Dracocephaletea aucheri cl. nov. hoc loco 1-1 Physoptychio gnaphalodis-Brometalia tomentosi ord. nov. hoc loco Syn.: Didymophysetea aucheri Klein & Lacoste 2001 nom. inval. (Art. 3b). Typus: Elymo longearistati-Astragalion macrosemii all. nov. Typus: Didymophysetalia aucheri ord. nov. (see below) (see below) Character species: Didymophysa aucheri, Dracocephalum auc- Character species: Bromus tomentosus, Physoptychis gnaphal- heri odes, Ziziphora clinopodioides Differential species (against Heldreichietea and Thlaspietea ro- Differential species: Poa araratica tundifolii): Alopecurus textilis This order covers the high-alpine and lower subnival This class comprises open plant communities on unsta- scree vegetation of Alborz and NW Iran and includes ble or stable screes in the alpine and subnival–nival zones mobile and stabilized scree grounds. Bromus tomentosus,

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Table 2. Abbreviated synoptic table of 18 alliances of scree vegetation in ﬁve regions of Europe and W Asia. C, character species; D, differential species. Only species with higher constancy are shown. For a longer version of this table, data sources and details on the calculation of constancy values, see Appendix S1.

Region Alps Balkans Caucasus Anatolia Iran

Column number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Number of releves 219 265 62 273 289 34 89 7 24 22 12 156 114 143 157 69 63 23

Widespread species Oxyria digyna 6 . 19 1 43 . . 43 5 90 . 1 1 4 1 13 . 4 Arabis alpina agg. 42 17 32 20 16 6 32 . . . . 11 32 4 14 10 . 26 (incl. A. caucasica) Cerastium cerastoides . . 3 3 2 . . . 10 43 . . . . 25 . . . Draba siliquosa . ..1....27...... 154 C Thlaspietea rotundifolii Cerastium uniflorum 1.66675...... Linaria alpina 51 25 34 2 18 12 30 ...... Poa cenisia s.lat. 2 3 . . . . 24 86 . . . . 1 10 3 . . . Ranunculus alpestris 23 7 5 64 7 6 ...... Rumex scutatus 22 49 . . 1 65 36 ...... Saxifraga oppositifolia 6 .79893...... Silene vulgaris subsp. 18 47 . 1 14 56 34 ...... glareosa et prostrata Achillea atrata 40 7 16 45 7 12 ...... Campanula cochleariifolia 37 46 15 6 1 18 ...... Pritzelago alpina 64 12 42 51 4 15 ...... Ranunculus montanus agg. 20 17 5 25 1 21 6 57 10 35 15 ...... Moehringia ciliata 53 18 3 36 . 6 10 ...... Poa minor 43 17 23 18 . 18 4 ...... Thlaspi rotundifolium 466.1 .9...... Adenostyles glabra 26 43 . 5 . 15 ...... Petasites paradoxus 140...182...... Saxifraga rudolphiana . .55. 5...... Salix retusa 7 1 11 53 24 3 4 ...... Poa laxa 1 .5.44...... Saxifraga androsacea 10.404218...... Saxifraga bryoides 1 .29376..14...... Achnatherum calamagrostis .4.. .262...... Epilobium dodonaei . ... .12...59...... Cardamine glauca ...... 40 43 ...... Saxifraga pedemontana ...... 57...... subsp. cymosa D Alps Viola biflora 40 22 . 30 3 21 ...... Minuartia gerardii 1114511103...... Saxifraga moschata 1114511213...... Persicaria vivipara 1523269263...... Silene acaulis subsp. exscapa . .15263...... Leucanthemopsis alpina 3.15356...... D Alps and Balkans Juncus trifidus 1 . . 1 30 . . 100 ...... Arenaria biflora . 23. 3.286...... Festuca violacea agg. 5 2 2 4 36 . . 71 ...... D Alps–Balkans–Caucasus Carex atrata agg. 1 . 11 42 3 3 . . . 39 15 ...... Phleum alpinum agg. 2 . 2 2 34 . . . . 45 ...... Dicranoweisia crispula ....5 .57.2070...... Dryopteris filix-mas . 1 . . 2 3 . 43 . 15 90 ...... Poa alpina s.lat. 35 14 69 66 61 3 . . . 38 15 ...... (incl. P. badensis)

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Table 2. (Continued). Region Alps Balkans Caucasus Anatolia Iran

Column number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Number of releves 219 265 62 273 289 34 89 7 24 22 12 156 114 143 157 69 63 23

DBalkans Senecio glaberrimus ...... 71...... Doronicum columnae . 1 . . . . 15 57 ...... C and D scree vegetation of Caucasus Sedum tenellum ...... 27 70 70 . . 2 6 6 . . Carum caucasicum ...... 5040.. . . .7.. Alopecurus ponticus ...... 552...... Veronica minuta ...... 5052...... Matricaria caucasica ...... 2378...... Murbeckiella huetii ...... 15 45 70 ...... Alopecurus dasyanthus ...... 45...... 6.. Potentilla gelida ...... 45...... Chaerophyllum humile ...... 63...... Corydalis alpestris ...... 45...... Dentaria bipinnata ...... 43...... Hyalopoa pontica ...... 23 52 15 ...... Lascuraea saxicola ...... 51050...... Common scree species Caucasus–Iran Saxifraga sibirica ...... 15 63 30 . . . . 26 . . Lamium tomentosum ...... 305...... 35 Senecio taraxacifolius ...... 27.. . . .48.. C and D Heldreichietea Heldreichia spp...... 17 33 19 31 . . . Euphorbia herniariifolia ...... 62 . 45 31 . . . Bunium microcarpum ...... 43271320... subsp. microcarpum Elymus tauri ...... 41 13 12 . . . Fritillaria crassifolia ...... 57...... Scrophularia myriophylla ...... 50. .... Anthriscus kotschyi ...... 429... Arenaria balansae ...... 4848... Lamium eriocephalum ...... 2550... C and D Didymophyso-Dracocephaletea Didymophysa aucheri ...... 32.96 Dracocephalum aucheri ...... 353248 Bromus tomentosus ...... 707813 Ziziphora clinopodioides ...... 4925. Poa araratica ...... 49374 Erigeron caucasicus ...... 62.. Festuca alaica ...... 7534 Astragalus macrosemius ...... 57. Elymus longearistatus ...... 60. Galium aucheri ...... 52

Physoptychis gnaphalodes and Ziziphora clinopodioides prefer NW Iran, Zagros and Alborz (Rechinger 1968). Ziziphora scree fields but can also be found scattered and with low clinopodioides is an Irano-Turanian element with nine frequency in alpine thorn cushion grasslands. Bromus subspecies in the Flora Iranica territory, where some of tomentosus plays an important role in plant communities them reach the alpine and subnival screes (e.g. subsp. el- of this order, reaching especially high cover in almost bursensis, filicaulis and pseudodasyantha), but most of them stabilized screes. It is distributed in Transcaucasia and the are rather common in sub-alpine and montane regions whole territory of Flora Iranica (Bor 1970). Physoptychis (Rechinger 1982b). Poa araratica is not exclusively con- gnaphalodes is a scree species of Transcaucasia, Anatolia, fined to scree fields but is also dominant in alpine thorn

102 10 Table 3. Typus releves. Association numbers are the same as in Table 1. For the association names, see Table 4. Letters after the association number refer to sub-associations. Iran of vegetation Scree

Relevenumber 123456789101112 Associationnumber 1 23a3b3c3d4 56a6b710 Turboveg releve number 1066 1072 1040 1035 1100 1088 950 760 832 718 888 916 Geographic coordinates 37°44′ 37°44′ 37°43′ 37°44′ 37°43′ 37°44′ 38°16′ 35°53′ 35°52′ 35°54′ 35°53′ 36°21′ 17.2″N 14.3″N 58.6″N 00.0″N 56.0″N 06.2″N 24.6″N 38.0″N 44.0″N 08.0″N 07.0″N 55.0″N 46°29′ 46°30′ 46°31′ 46°31′ 46°29′ 46°29′ 47°51′ 51°24′ 51°24′ 51°24′ 51°24′ 50°57′ 53.3″E 09.5″E 08.6″E 09.3″E 58.5″E 59.7″E 15.0″E 32.0″E 35.0″E 40.0″E 54.0″E 44.0″E

Nepeta menthoides 41...1...... Alopecurus aucheri 2...... Senecio taraxacifolius + 2...1...... PhD ThesisofJalilNorooziEshlaghi2013 Myosotis asiatica . + ...... Oxyria digyna .1...... Jurinella moschus ..1...... Potentilla porphyrantha ..2.+ .1..... Minuartia recurva subsp. oreina ..+ ...... Astragalus beckii ....1...... Draba bruniifolia subsp. bruniifolia ....2.+ ..... 103 Nepeta sahandica .....1......

o:10.1111/avsc.12031Doi: Thymus kotschyanus .....+ ...... Crepis frigida .....+ ...... Erysimum gelidum .....+ ...... Alopecurus dasyanthus ...... + ..... Tripleurospermum caucasicum .1....1..... Saxifraga sibirica .....++ ..... Erigeron caucasicus subsp.venustus .11+ 1 ++ ..... Sesleria phleoides ..2...... © + 03ItrainlAscainfrVgtto Science Vegetation for Association International 2013 Galium hyrcanicum .11...... Pedicularis caucasica ..+ . ++ ...... Minuartia glandulosa .1.+ ...... Koeleria eriostachya + .1.1...... Scutellaria glechomoides ...... 2.... Euphorbia aucheri ...... 21+ .. Nepeta racemosa ...... 33..

Cicer tragacanthoides ...... 3... Chapter 3 + ple eeainScience Vegetation Applied Leonurus cardiaca subsp. persicus ...... Polygonum serpyllaceum ...... 1.. Asperula glomerata subsp. bracteata ...... 1. Draba siliquosa ...... + .....

Astragalus macrosemius ...... 2. al. et Noroozi J. Elymus longearistatus ...... 121. Bromus tomentosus + 11121. .1.2. Ziziphora clinopodioides ..1121....1. Physoptychis gnaphalodes ...11...... PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

J. Noroozi et al. Scree vegetation of Iran ′ ′ N E ″ ″ cushion grasslands (Noroozi et al. 2010). It is distributed 21 57 ° ° from Anatolia and the Caucasus to the Hindu Kush (Bor 36 55.0 50 44.0 1970). Two alliances are distinguished within the order. One ′ ′ N E ″ ″ 53 24 2. ° ° alliance is distributed in Alborz and the other in Sahand 35 07.0 51 54.0 and Sabalan (NW Iran). ′ ′ N E ″ ″ 1-1-1 Erigerontion venusti all. nov. hoc loco (Appendix S4) 54 24 + ° ° 35 08.0 51 40.0 Typus: Dracocephalo aucheri-Brometum tomentosi ass. nov. (see below) ′ ′ N E ″ ″ Character taxa: Erigeron caucasicus subsp. venustus, Nepeta 52 24 ° ° menthoides, Senecio taraxacifolius, Tripleurospermum caucasi- 35 44.0 51 35.0 cum var. caucasicum, Tripleurospermum caucasicum var. melanolepis ′ ′ E N ″ ″ 24 53 ....1 ....1 ° ° Differential species: Alopecurus textilis, Draba bruniifolia 51 32.0 35 38.0 subsp. bruniifolia, Erysimum gelidum, Festuca alaica, Galium hyrcanicum, Koeleria eriostachya, Minuartia glandulosa, Pedicu- ′ ′ N E ″ ″ laris caucasica, Saxifraga sibirica, Sesleria phleoides 16 51 + ° ° This alliance covers the scree vegetation of Sahand and 47 38 24.6 15.0 Sabalan. Erigeron caucasicum subsp. venustus is an alpine– subnival species distributed from NE and E Anatolia to N ′ ′ E N ″ ″ 29 44 ...... ++ ° ° Iran (Grierson 1975; Rechinger 1982a). It is common in 46 59.7 37 06.2 Sahand and Sabalan but very rare in Alborz. Nepeta menthoides is recorded from NE Iraq and NW Iran (Rechinger ′ ′ E N ″ ″ 1982b). Senecio taraxacifolius is distributed in NE Anatolia, 29 43 + ° ° Caucasus and NW Iran (Nordenstam 1989). In Iran, it is 46 58.5 37 56.0 restricted to the high-alpine–subnival scree habitats of Sa- hand and Sabalan. Tripleurospermum caucasicum occurs on ′ ′ N E ″ ″ 44 31 .21. .. ° ° the Balkan Peninsula, in Lebanon, Anatolia, NE Iraq, the 37 00.0 46 09.3 Caucasus and NW Iran (Podlech 1986). There are two varieties of this species in Iran, var. caucasicum and var. ′ ′ E N – ″ ″ melanolepis, which are both distributed in the alpine sub- 31 43 .. .111....12 1...... 1. + ° ° nival zone of E Anatolia, the Caucasus and NW Iran 46 08.6 37 58.6 (Podlech 1986). The diagnostic species include several differential species that are equally common in thorn ′ ′ N E ″ ″ 44 30 + + + ° ° cushion grasslands. Festuca alaica is distributed from NW 37 14.3 46 09.5 Iran to the Hindu Kush and Central Asia (Alexeev 1979). Saxifraga sibirica, which also grows on rocky habitats, is a ′ ′ E N ″ ″ widespread species in Eurasia, occurring from mountains 29 44 ...... 1 ....11 ...... 1...... 1...... 1...... 1...... 1...... 111111...... ° ° of the Balkan Peninsula to Russia (including Siberia), in 46 53.3 17.2 the Caucasus, NW Iran and Central Asia (Schonbeck-€ Temesy 1967).

1-1-1-1 Nepetetum menthoidis ass. nov. hoc loco (Table 1: 1, beauverdiana Appendix S4, Photo S2)

subsp. Typus: releve 1 (ﬁeld nr. 1066) in Table 3. Character species (transgressive):

e number 1066 1072 1040 1035 1100 1088 950 760 832 718Nepeta 888menthoides 916 Differential species: Alopecurus aucheri, Carum caucasicum, (Continued). Sedum tenellum, Taraxacum crepidiforme nme 123456789101112 enumber This association is found on wet scree ground with high Dracocephalum aucheri Helichrysum psychrophilum Cerastium purpurascens Didymophysa aucheri Acantholimon demawendicum Arenaria insignis Veronica orientalis Linaria lineolata Campanula stevenii Festuca alaica Arabis caucasica Poa araratica Alopecurus textilis soitonme 3333456a6b710 1 23a3b3c3d4 Associationnumber Turboveg relev Geographic coordinates 37 Table 3. Relev cover of Nepeta menthoides, which is a large perennial herb

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Table 4. Syntaxonomic survey of the study area. which is also indicated by the presence of snowbed species

Class Didymophyso aucheri-Dracocephaletea aucheri cl. nov. such as Carum caucasicum, Alopecurus aucheri and Taraxacum Order 1 Physoptychio gnaphalodis-Brometalia tomentosi ord. nov. crepidiforme Alliance 1 Erigerontion venusti all. nov. Ass. 1 Nepetetum menthoidis ass. nov. Ass. 2 Senecionetum taraxacifolii ass. nov. 1-1-1-2 Senecionetum taraxacifolii ass. nov. hoc loco (Table 1: Ass. 3 Dracocephalo aucheri-Brometum tomentosi ass. nov. 2, Appendix S4, Photo S3) Ass. 4 Potentilletum porphyranthae ass. nov. Typus: releve 2 (ﬁeld nr. 1072) in Table 3. Alliance 2 Elymo longearistati-Astragalion macrosemii all. nov. Ass. 5 Scutellario glechomoidis-Euphorbietum aucheri ass. nov. Character species (transgressive): Senecio taraxacifolius Ass. 6 Cicero tragacanthoidis-Nepetetum racemosae ass. nov. Differential species: Myosotis asiatica, Oxyria digyna Ass. 7 Asperulo bracteatae-Brometum tomentosi ass. nov. This association is widespread in the high-alpine and Ass. 8 Senecioni iranici-Astragaletum macrosemii ass. prov. subnival belt of Sahand in N aspects with steep inclination Order 2 Didymophysetalia aucheri ord. nov. (ca. 30–45°). It occurs at altitudes from ca. 3300 m up to Alliance 3 Didymophysion aucheri all. nov. the highest peak, 3707 m a.s.l. The ground is covered with Ass. 9 Galietum aucheri Klein & Lacoste 2001 big and small mobile and stabilized stones. Senecio taraxacifo- Ass. 10 Dracocephaletum aucheri ass. nov. lius is the most dominant species of this association, and its yellow ﬂowers make the stands showy and attractive in July. Myosotis asiatica and Oxyria digyna are widespread scree species in the northern hemisphere (Meusel et al. 1964).

1-1-1-3 Dracocephalo aucheri-Brometum tomentosi ass. nov. hoc loco (Table 1: 3, Appendix S4, Photos S4, 5, 6)

Typus: releve 5 (field nr. 1100) in Table 3. Central association of the alliance. Character species (only present in certain sub- associations): Astragalus beckii, Nepeta sahandica This is one of the most widespread associations of the highest altitudes of Sahand, growing in SW, W and NW aspects with ca. 20–30° inclinations and an altitudinal range of ca. 3500–3707 m a.s.l. Four sub-associations can be recognized: 1 Dracocephalo-Brometum jurinelletosum moschi subass. nov. hoc loco [Typus: releve 3 (field nr. 1040) in Table 3] (Photo S4) with Jurinella moschus, Potentilla porphyrantha and Minuartia recurva subsp. oreina as differential taxa. This community is recorded from W slopes near ridges at Fig. 2. Detrended correspondence analysis diagram of the sample plots. altitudes of around 3500–3550 m a.s.l. The ground is Association numbers are the same as in Table 1. 1: Nepetetum often covered with small-grained scree. J. moschus is typi- menthoidis,2:Senecionetum taraxacifolii,3:Dracocephalo aucheri- cal for scree vegetation near the top of mounds and Brometum tomentosi,4:Potentilletum porphyranthae,5:Scutellario ridges of the alpine and subnival areas of Anatolia, glechomoidis-Euphorbietum aucheri,6:Cicero tragacanthoidis-Nepetetum Transcaucasia, NE Iraq and NW Iran (Rechinger 1979). racemosae,7:Asperulo bracteatae-Brometum tomentosi,8:Senecioni The alliance Jurinellion moschi Parolly 1995 was described iranici-Astragaletum macrosemii,9:Galietum aucheri,10:Dracocephaletum aucheri. Left circle: Erigerontion venusti (Ass. 1–4), right circle: Elymo- from scree habitats of the Taurus Mts. in Central Anato- Astragalion macrosemii (Ass. 5–8), upper circle: Didymophysion aucheri lia, which is classified within the Heldreichietalia Quezel (Ass. 9–10). ex Parolly 1995 and Heldreichietea Quezel ex Parolly 1995 (Parolly 1995). of up to 60 cm. The stands are located in E and NE aspects 2 Dracocephalo-Brometum minuartietosum glandulosae between alpine thorn cushion grasslands and steep scree subass. nov. hoc loco [Typus: releve 4 (field nr. 1035) in slopes. The high moisture of the soil is due to heavy snow Table 3] (Photo S5) with Minuartia glandulosa as differen- accumulation during winter. In the DCA, the association is tial species. This unit is recorded from W aspects with ca. clearly separated at the left side of the diagram (Fig. 2). It is 30° inclination at ca. 3510–3540 m a.s.l. The ground is a transition between scree and snowbed communities, covered with big stones (30–50 cm) and smaller scree

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Fig. 3. Box plots for altitude, ‘southness’ of aspect, ‘westness’ of aspect, inclination, species number per plot and vegetation cover for all three alliances and ten associations. The boxes represent the interquartile range (IQR), horizontal lines the median values and points the outliers (>1.5 IQR outside the box). Alliance 1 (Ass. 1–4): Erigerontion venusti,Alliance2(Ass.5–8): Elymo-Astragalion macrosemii, Alliance 3 (Ass. 9–10): Didymophysion aucheri. The association numbers are the same as in Table 1 and Fig. 2. For the association names, see Table 4. material. Minuartia glandulosa is a cushion plant and more bruniifolia, Alyssum condensatum and Alopecurus textilis. This dominant in the alpine thorn cushion grasslands of unit has a higher vegetation cover, and Bromus tomentosus, Sahand. It is distributed in E Anatolia, Armenia, Republic Poa araratica and Ziziphora clinopodioides are more dominant of Azerbaijan and NW Iran (Rechinger 1988). than in the other three sub-associations. Astragalus beckii is 3 Dracocephalo-Brometum typicum subass. nov. hoc loco distributed in Alborz and NW Iran as well as in Talysh [Typus = typus of the association] (Photo S6) is more (Podlech et al. 2010). Draba bruniifolia subsp. bruniifolia widespread than the two previous sub-associations and and Alyssum condensatum are common dominant species in occurs at the highest elevations of Sahand (ca. 3600– alpine thorn cushion grassland. The ﬁrst is distributed in 3707 m a.s.l.) in SW, W and NW aspects. The character NW Iran, SE Anatolia and NE Iraq (Hedge 1968), and the species Astragalus beckii seemstoberestrictedtothissub- second in Lebanon, Syria, Anatolia and NE Iraq (Rechinger association. Differential taxa are Draba bruniifolia subsp. 1968).

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The plant cover and species richness are very low. Potentilla porphyrantha is a narrowly ranged species of the subnival zones of S Armenia and NW Iran (see distribution map in Noroozi et al. 2011). Potentilla argaea is distributed in Ana- tolia and NW Iran (Schiman-Czeika 1969). Alopecurus dasyanthus occurs in NE Anatolia, the Caucasus and NW Iran (Bor 1970).

1-1-2 Elymo longearistati-Astragalion macrosemii all. nov. hoc loco (Appendix S5)

Typus: Asperulo bracteatae-Brometum tomentosi ass. nov. (see below) Fig. 4. Mean proportional cover of life forms over all releves of each Character species: Euphorbia aucheri, Scutellaria glechomoides association. Ass. 1–4: Erigerontion venusti,Ass.5–8: Elymo-Astragalion Differential species: Astragalus macrosemius, Elymus longeari- macrosemii, Ass. 9–10: Didymophysion aucheri. The association numbers status are the same as in Table 1 and Fig. 2. For the association names, see This alliance includes the high-alpine to lower subnival Table 4. Non-Cush Cham: non-cushion chamaephyte, Cush Cham: cushion scree vegetation of Central Alborz. Euphorbia aucheri and chamaephyte, Tall-Hemi: tall hemicryptophyte (>ca. 30 cm), Small-Hemi: small hemicryptophyte (

107 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

J. Noroozi et al. Scree vegetation of Iran the Hindu Kush (Rechinger & Schiman-Czeika 1964). Scu- Central association of the alliance. tellaria glechomoides is restricted to Central Alborz (Rechin- This association mostly occurs on SW slopes mostly ger 1982b). The community is known from 3230 to of 20–35° inclination at the highest altitudes of the Tu- 3800 m a.s.l. on S slopes but may extend up to the subni- chal (3800–3960 m a.s.l.) N of Tehran. The structure val–nival zone. and ecology of this association is very close to the Dra- cocephalo-Brometum tomentosi that covers the highest altitudes of Sahand, mostly in W aspects. The most 1-1-2-2 Cicero tragacanthoidis-Nepetetum racemosae ass. nov. remarkable physiognomic difference between the two hoc loco (Table 1: 6, Appendix S5, Photo S9A, 10,11) associations is the absence of thorn cushions in Draco- Typus: releve 9 (field nr. 832) in Table 3. cephalo-Brometum. In the DCA diagram these two associ- Character species: Cicer tragacanthoides, Leonurus cardiaca ations are located very close to each other near the subsp. persicus, Nepeta racemosa centre of the diagram. This association occurs on recently stabilized scree fields, Three provisional sub-associations have been distin- and sometimes is surrounding stands of the Scutellario-Eu- guished: phorbietum (see Photo S9a). According to field observations 1 The typical sub-association with Asperula glomerata of the first author, it is the most frequent association in the subsp. bracteata as differential species (Photo S12). A. glom- alpine scree slopes of Central Alborz. The altitudinal range erata is an altitudinally widespread species ranging from ca. of the community is ca. 3200–3730 m a.s.l., and it usually 700 to 4200 m a.s.l., with many subspecies. A. glomerata covers S slopes with inclinations of 15–45°. Nepeta racemosa subsp. bracteata is a strongly compact cushion restricted to is distributed in Alborz, with a single record from SE Tur- Central Alborz (Ehrendorfer et al. 2005) and concentrated key near the border with Iran (Rechinger 1982b). Cicer tra- in high-alpine and subnival scree habitats. gacanthoides, which has a tragacanthic habit, is restricted to 2 On N aspects with moderate slope, stabilized scree and the alpine zone of Iran (van der Maesen 1979). It crawls more developed soil, transitions to snowbed communities over the scree and thus stabilizes it (Photo S3). Leonurus can be found that are differentiated by Ranunculus crymo- cardiaca subsp. persicus is endemic in Central Alborz (Rech- philus and Polygonum serpyllaceum. Both of these are charac- inger 1982b). ter species of snowbed communities (Noroozi et al. 2010). Besides the typical sub-association (1) Cicero tragacan- 3 On steep slopes with large volcanic stony ground and thoidis-Nepetetum racemosae typicum subass. nov. hoc loco undeveloped soils on N and W slopes, Asperula glomerata [Typus = typus of the association] (Photo S10), a deviat- disappears and the dominance of Dracocephalum aucheri ing subunit (2) Cicero tragacanthoidis-Nepetetum racemosae increases (Photo S13). The ecology and physiognomy of polygonetosum serpyllacei subass. nov. hoc loco [Typus: this sub-association is very similar to Dracocephalo-Brome- releve 10 (field nr. 718) in Table 3] (Photo S11), can be tum minuartietosum glandulosae of Sahand (compare Photos distinguished on NE slopes with steep inclination (ca. 40 S5 and S13). –45°), where the ground is covered with coarse scree and boulders (ca. 0.5–1.0-m diameter). This sub-associa- 1-1-2-4 Senecioni iranici-Astragaletum macrosemii ass. prov. tion is differentiated by Polygonum serpyllaceum, a charac- (Table 1: 8, Appendix S5, Photo S14) ter species of snowbed communities (Taraxaco brevirostre- Polygonion serpyllacei Noroozi et al. 2010). Its presence in Character species: Senecio iranicus this community is due to long snow cover and high soil Differential species: Artemisia melanolepis, Carex oreophila, moisture. Nepeta racemosa has high cover in this sub- Catabrosella parviflora, Cerastium purpurascens var. elbursensis, association but the other two character species of the Draba siliquosa, Erysimum elbrusense, Potentilla argyroloma, association are absent. There is some ecological similarity Veronica biloba between this sub-association and the Nepetetum menthoi- This community reaches the highest elevation within dis of Sahand. In both communities, there are snowbed this alliance and probably harbours the highest occur- species, and both occur in NE and E aspects. The two rences of the spiny cushions in in the study area. In spite Nepeta species (N. racemosa and N. menthoides) play a sim- of the high altitude of the stands (ca. 4200–4230 m a.s.l.), ilar ecological role in the two different geographic finer material with appropriate moisture, moderate incli- regions. nation (ca. 10–15°) and stabilized scree make this community one of the richest of the class (ca. 14 species on 25 m²) with high vegetation cover (ca. 70%). This com- 1-1-2-3 Asperulo bracteatae-Brometum tomentosi ass. nov. hoc munity is located on the S slopes of Mt. Damavand. Senecio loco (Table 1: 7, Appendix S5, Photo S12, 13) iranicus is an annual and local endemic species for Damav- Typus: releve 11 (field nr. 888) in Table 3. and (Nordenstam 1989) and recorded only from this com-

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Scree vegetation of Iran J. Noroozi et al. munity. Artemisia melanolepis is an endemic species in Character species: Galium aucheri, Senecio vulcanicus, Veron- the subnival zone of NW and N Iran (Podlech 1986). Carex ica paederotae oreophila occurs in the Caucasus, Turkey, Iraq and Iran Differential species: Asperula glomerata subsp. bracteata, La- (Kukkonen 1998). Catabrosella parviflora is a dominant mium tomentosum, Pseudocamelina glaucophylla species of snowbed and thorn cushion grasslands of This association is concentrated at altitudes between Iranian high-alpine regions (Noroozi et al. 2010). Draba 3800–4200 m a.s.l., with ca. 20–35° inclination. It was siliquosa is a widespread species in European and Cauca- recorded by Klein & Lacoste (2001) from Alamkuh and sian alpine and subnival zones, and Damavand is the east- Azadkuh. In the DCA diagram, it is located in the centre of ernmost limit of this species (see distribution map in the alliance (Fig. 2). Galium aucheri, Senecio vulcanicus and Noroozi et al. 2011). Potentilla argyroloma is an endemic Veronica paederotae are subnival–nival species and restricted species of Iran and occurs in Zagros and Alborz (Schiman- to Alborz (see distribution maps in Fischer 1981; Czeika 1969). Veronica biloba is an annual species with a Nordenstam 1989; Ehrendorfer et al. 2005; Noroozi et al. wide altitudinal (ca. 1200–4220 m a.s.l.) and geographical 2011). The main distribution of Lamium tomentosum is distribution from the Caucasus and W Asia to Central Caucasus and Transcaucasus mountains, with disjunct Asia, Mongolia and the Himalaya (Fischer 1981). Ceras- distribution in Central Alborz (Mennema 1989; see tium purpurascens var. elbursensis and Erysimum elbrusense distribution map in Noroozi et al. 2011). Pseudocamelina are diagnostic taxa of the Didymophysion aucheri (see glaucophylla is an endemic species of Iran but widespread in below). the alpine zone of Zagros, NW Iran and Alborz mountains (Rechinger 1968). Klein & Lacoste (2001) indicated also Achillea aucheri and Scutellaria glechomoides as character 1-2 Didymophysetalia aucheri ord. nov. hoc loco species of this association. According to our data, however, Typus: Didymophysion aucheri all. nov. (see below) these two species clearly have their optimum in other Character species (transgressive): Didymophysa aucheri, Dra- communities. cocephalum aucheri Differential species: Erysimum elbrusense 1-2-1-2 Dracocephaletum aucheri ass. nov. hoc loco (Table 1: This order is monotypic, with a single alliance so far (but 10, Appendix S6, Photo S16, 17) see Potentilletum porphyranthae above). Erysimum elbrusense is common in Alborz but is also known with a few records Typus: releve 12 (field nr. 916) in Table 3. from NW Iran (Polatschek 2011). Central association of the alliance. This association represents the upper limit of vascular plants in Central Alborz. It is the highest plant community 1-2-1 Didymophysion aucheri all. nov. hoc loco (Appendix S6) of the Iranian mountains, concentrated between 4300 and Typus: Dracocephaletum aucheri ass. nov. (see below) 4600 m a.s.l., but in some special places reaches even Character species: Achillea aucheri, Cerastium purpurascens higher altitudes. The ground is completely covered with var. elbursensis scree. This alliance includes open communities at the upper Two subunits can be distinguished which, due to the limit of vascular plant species in the subnival–nival zone of small number of releves, are treated as provisional sub- Central Alborz. Very low cover–abundance of grasses, associations. With more material, however, the first unit absence of thorn cushions, low species richness, sparse might become a good association, with Achillea aucheri and plant cover and a high percentage of open scree are the Veronica aucheri as character species. main features of this alliance. Achillea aucheri isatruesub- 1 Dracocephaletum aucheri achilleetosum aucheri subass. prov. nival–nival species in Central Alborz, with a few records (Photo S16) with the diagnostic species Achillea aucheri and from NW Iran (Huber-Morath 1986). Cerastium purpuras- Veronica aucheri. This community grows at very high alti- cens is an alpine and subnival–nival species of Anatolia, the tudes of Damavand on S slopes from 4300 to 4800 m a.s.l., Caucasus, NE Iraq and Central Alborz. In Iran it is confined and future studies may lead to the conclusion that it to the subnival–nival zone of Central Alborz, a taxon should be treated as a separate association. Veronica aucheri known as var. elbursensis (Moschl€ 1988). is a true subnival–nival species and endemic for Alborz (Fischer 1981). It is the most common species at altitudes from 4500 to 4800 m a.s.l. of Damavand, and has the high- 1-2-1-1 Galietum aucheri Klein & Lacoste 2001 [‘1999’] est altitudinal record for vascular plants in Iran (4820 m (Table 1: 9, Appendix S6, Photo S15) a.s.l.). In one plot of this association (4799 m a.s.l.), Alope- Typus: releve 99 in Klein & Lacoste 2001 (see Appendix curus himalaicus was found, which has an extremely dis- S6, releve no. 1170) junct distribution in Bulgaria, Iran and the W Hindu Kush

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–Himalaya system (Noroozi et al. 2011). This is probably to be similar to that on Mediterranean mountains of S Eur- the same location from which the species was previously ope, where species richness has decreased during the past recorded (Bor 1970), and so far the only known popula- particularly warm decade (Pauli et al. 2012). Climate tion of this species in Iran. change models predict increasing temperature and 2 Dracocephaletum aucheri typicum (Photo S17) with Arabis decreasing precipitation for our study region during the caucasica and Potentilla argyroloma as differential species. It 21st century (Christensen et al. 2007; Zarghami et al. is recorded from the nival zone of Mt. Alamkuh at altitudes 2011). Hence, we have to presume a very high risk of bio- from 4350 to 4450 m a.s.l. on gentle to steep slopes with- diversity losses in the hig-altitude vegetation of N and NW out special aspect preference. Iran. Detailed knowledge of these plant assemblages appears to be essential for tracing the expected changes in their species composition and the possible shrinkage of Concluding remarks cryophilic, narrowly distributed species. Floristically defined vegetation types are suitable reference entities for ecological research, bioindication and nature Acknowledgements conservation (Dengler et al. 2008). Our study extends We thank J. Dengler and J.-P. Theurillat for valuable com- knowledge of alpine scree vegetation into a previously ments on a former version of the manuscript. M. Djamali is almost unknown area, and therefore represents an impor- acknowledged for providing information on the geology tant building block for a future synopsis of the scree vege- and climate of the study area. We thank N. Sauberer tation of the northern hemisphere. The alpine and (Gramineae), F. Ehrendorfer (Rubiaceae), D. Podlech subnival scree communities in our study area show a high (Astragalus), A. Polatschek (Erysimum), B. Wallnofer€ degree of local endemism, and many endemic taxa of Iran (Carex) and H. Scholtz (Festuca and Bromus) for determina- are restricted to high altitudes (Klein 1991; Frey et al. tion of certain critical taxa. This work was partly supported 1999; Noroozi et al. 2008, 2011). We therefore consider by the GLORIA co-ordination at the University of Vienna our study as an important assessment tool for nature con- and the Austrian Academy of Sciences (www.gloria.ac.at). servation that will improve knowledge of phytodiversity in alpine habitats of Iran. It is also a basis for monitoring of future changes and for assessing endangered vegetation References types. Agakhanjanz, O. & Breckle, S.-W. 1995. Origin and evolution of The direct human impact on the described scree com- the mountain flora in middle Asia and neighboring moun- munities is not very strong due to their low productivity. tain regions. In: Chapin, F.S. & Korner,€ C. (eds.) Arctic and However, there are large herds grazing neighbouring alpine biodiversity: patterns, causes and ecosystem consequences,pp. alpine grasslands. Therefore, in spite of the low attraction 63–80. Ecological Studies 113, Springer, Berlin, DE. for sheep and goats to feed on scree vegetation, effects of Akhani, H., Mahdavi, P., Noroozi, J. & Zarrinpour, V. 2013. Veg- pastoralism should not be neglected, especially the impact etation pattern of Irano-Turanian steppes along a 3000 m of trampling on unstable scree habitats. As the herds in altitudinal gradient in the Alborz Moutains (Iran). Folia Geo- these mountains are usually very large, the substrate botanica. doi:10.1007/s12224-012-9147-8. movement is strongly intensified by trampling, plants are Alexeev, E. 1979. Genus Festuca L. Florae Iranicae et territorium – uprooted and thus the vegetation is sometimes completely confinium. Novitates systematicae Plantarum vascularium 16: 7 destroyed. The most important threat to the flora and veg- 35. etation of the study habitats, however, may be the acceler- Behrouzi, A., Amini Fazl, A., Amini Azar, A. & Emami, M.H. 1997. Geological map of Bostan Abad (1: 100,000). Geological ation of climate warming. The high rate of narrowly Survey of Iran, Tehran, IR. distributed species in the alpine and subnival–nival zones Bor, N.L. 1970. Gramineae. In: Rechinger, K.H. (ed.) Flora ira- and the absence of alternative low-temperature habitats in nica, 70,pp.1–573. Akademische Druck u. Verlagsanstalt, Iranian mountains increase this threat (Noroozi et al. Graz, AT. 2011). The highest growing vascular plant in Iran was Buhse, F. 1899a. Flora des Alburs und der Kaspischen Sudk€ uste.€ found at ca. 4800 m a.s.l. on Damavand, and only three Bisherige Forschungsergebnisse aus diesem Gebiet. Arbeiten mountain peaks exceed the upper limit of vascular plants des Naturforscher-Vereins zu Riga. Neue Folge.8.Heft. (Damavand and Alamkuh in Central Alborz and Sabalan Buhse, F 1899b. Reisebemerkungen aus dem ostlichen€ Albu- in NW Iran). Therefore, global warming will threaten the rsgebirge in Persien. Bulletin de la SocieteImperiale des Natural- very fragmented and small-scale habitats of the cryophil- istes de Moscou 34: 363–383. ous flora and vegetation of the Iranian mountains (Noroozi Christensen, J. H., Hewitson, B., Busuioc, A., Chen, A., Gao, X., et al. 2011). A decline of high-altitude habitats through cli- Held, I., Jones, R., Kolli, R. K., Kwon, W.-T., Laprise, R., mate warming and reduced water availability is expected Magana~ Rueda, V., Mearns, L., Menendez, C. G., Rais€ anen,€

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Willner, W. 2011. Unambiguous assignment of releves to vegeta- Appendix S1. Synoptic table of the scree vegetation in tion units: the example of the Festuco-Brometea and Trifolio- ﬁve regions of Europe and western Asia. Geranietea sanguinei. Tuexenia 31: 271–282. Appendix S2. The mean and SD of the environmental Willner, W., Tichy, L. & Chytry, M. 2009. Effects of different ﬁdelity measures and contexts on the determination of diag- and vegetation parameters in all associations. nostic species. Journal of Vegetation Science 20: 130–137. Appendix S3. Synoptic table showing the total cover and Wright, H. 1962. Pleistocene glaciation in Kurdistan. Eiszeitalter diagnostic value of species in all vegetation units. – und Gegenwart 12: 131 164. Appendix S4. Phytosociological table of the alliance Eri- Zarghami, M., Abdi, A., Babaeian, I., Hassanzadeh, Y. & Kanani, gerontion venusti. R. 2011. Impacts of climate change on runoffs in East Azer- Appendix S5. baijan, Iran. Global and Planetary Change 78: 137–146. Phytosociological table of the alliance Zarre, S., Massoumi, A.A. & Podlech, D. 2008. Papilionaceae V, Elymo longearistati-Astragalion macrosemii. Astragalus III. In: Rechinger, K.H. (ed.) Flora Iranica, 177,pp. Appendix S6. Phytosociological table of the alliance 1–124. Naturhistorisches Museum Wien, Wien, AT. Didymophysion aucheri. Zohary, M. 1973. Geobotanical foundations of the Middle East,Vol. Photos S1–S17. Typical stands of the associations 2. Gustav Fischer, Stuttgart, DE. described in this study.

Supporting Information Additional supporting information may be found in the online version of this article:

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Appendix 2a. The mean and standard deviation of the environmental and vegetation parameters in all associations. Different superscript letters after the mean of variables indicate significant differences between the associations (p < 0.05; Games-Howell post hoc test). A “-“ indicates that the post hoc test was not possible because of zero variance. The p-value of the ANOVA was highly significant for all variables (p < 0.001). Note that for ass. 9 (plots of Klein & Lacoste 1999) no data of scree, bare ground and solid rock covers are available. Ass. 1-4: Erigerontion venusti, Ass. 5-8: Elymo-Astragalion macrosemii, Ass. 9-10 : Didymophysion aucheri. For association names see Fig. 2 or Table 4.

No. of Relevé Altitude Southing Westing Inclination Species Shannon Evenness Vegetation Scree Bareground Solidrock Association relevés size (m2) (m a.s.l.) (degree) (degree) (degree) richness Index Index cover (%) cover (%) cover (%) cover (%) Mean 25 3356a 104a 166a 37a 9a 1.28a 0.46a 61a 34a 6a 0a 1 8 SD 0 4 11 11 3 4 0.46 0.09 14 14 3 0 Mean 100 3420a 137b 109b 38a 10a 1.80ab 0.68b 34b 40a 12a 15b 2 19 SD 0 107 25 46 5 4 0.75 0.2 6 12 8 14 Mean 94 3599b 123ab 49c 28b 11a 2.10bc 0.78b 22c 64b 6a 7bc 3 35 SD 21 69 38 48 9 3 0.31 0.13 11 17 4 11 Mean 100 4310c 180- 90- 14c 6ab 1.70ac 0.98c 6d 13a 6a 74 4 7 SD 0 62 0 0 2 3 0.53 0.01 4 7 2 9 Mean 25 3556abd 23c 67bc 19bc 2b 0.75a 0.90bc 10de 89c 1ab 0a 5 6 SD 0 246 20 20 7 1 0.21 0.15 3 4 2 0 Mean 25 3574b 55c 127b 35ab 7a 1.38a 0.59ab 42ab 45a 1bc 8ab 6 21 SD 0 151 55 36 9 1 0.33 0.13 13 17 3 13 Mean 41 3906d 63c 77bc 24b 6a 1.37ac 0.69b 34b 62b 3ac 2ac 7 33 SD 31 48 57 40 11 2 0.41 0.15 11 13 3 3 Mean 25 4218c 0- 90- 13c 14a 1.98ac 0.58ab 68abcef 17a 8ac 7ab 8 3 SD 0 4 0 0 3 2 0.23 0.01 12 3 6 3 Mean 114 3900d 58abc 84b 26b 7a 1.89ac 0.98c 7dfg - - - 9 12 SD 152 166 75 22 9 2 0.3 0.01 2 - - - Mean 65 4429c 45c 69bc 16bc 6a 1.44a 0.78bc 12cdg 60ab 3ac 21ab 10 11 SD 40 135 39 57 10 3 0.51 0.18 10 20 3 24 Mean 66 3754 88 86 28 8 1.63 0.73 28 49 5 10 Total 155 SD 56 319 61 50 11 4 0.55 0.19 18 25 5 19

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Appendix 2b. The mean and standard deviation of the environmental and vegetation parameters in the three alliances. Different superscript letters after the mean of variables indicate significant differences between the alliances (p < 0.05; Games-Howell post hoc test). The p-value of the ANOVA is given at the bottom of the table. All. 1: Erigerontion venusti, All. 2: Elymo-Astragalion macrosemii, All. 3: Didymophysion aucheri.

No. of Relevé Altitude Southing Westing Inclination Species Shannon Evenness Vegetation Scree Bareground Solidrock Alliances relevés size (m2) (m a.s.l.) (degree) (degree) (degree) richness Index Index cover (%) cover (%) cover (%) cover (%) Mean 88 3594a 130a 83 31a 10a 1.88a 0.73a 28a 48a 8a 15a 1 69 SD 28 271 36 58 10 3 0.56 0.19 17 22 6 23 Mean 33 3777b 54b 93 27ab 6b 1.35b 0.67a 36b 57a 2b 4b 2 63 SD 24 225 54 43 12 3 0.42 0.16 16 21 3 8 Mean 91 4153c 52b 77 21b 6b 1.67a 0.89b 9c 28b 1b 10ab 3 23 SD 114 308 60 42 10 2 0.46 0.16 7 33 2 19 Mean 66 3752 88 86 28 8 1.63 0.73 28 49 5 10 Total 155 SD 56 320 61 50 11 4 0.55 0.19 18 25 5 19 p (ANOVA) - - 0 0 0.329 0.002 0 0 0 0 0 0 0.001

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Appendix S3. Synoptic table showing the total cover and diagnostic value of species in all vegetation units. Values are total cover (TC), i.e., average cover taking also into account zero values. Species cover in individual relevés was approximated by taking the average cover corresponding to each value of the "Braun-Blanquet scale". TC values of higher syntaxa are averages of the respective values in the subordinated units. The values in ass. 8 (given in red) were downweighted by a factor of 10 because of the low number of relevés and the marginal position of the association in the alliance. Fidelity of species was calculated using the Total Cover Ratio (TCR), i.e., the ratio between the highest and second highest TC among the units of the same rank within a higher syntaxon. TC values of diagnostic species are shaded in yellow (if TCR > 2) or green (if TCR > 10). See text for details on the determination of diagnostic species. C: character species, tC: transgressive character species, D: differential species. Association numbers are the same as in Table 1. For the association names, see Table 4.

Associations Alliances Orders Order P-B P-B P-B P-B P-B P-B P-B P-B Da Da P-B P-B Da P-B Da Alliance Ev Ev Ev Ev E-Am E-Am E-Am E-Am Da Da Ev E-Am Da Association 12345678910 Number of relevés 8 19 35 7 6 21 33 3 12 11 69 63 23 132 23

C & D Ass. 1-4 Nepeta menthoides (tC Ass. 1) 49.32.80.1...... 13.1..6.5. Alopecurus aucheri 6.81.5...... 2.1..1.0. Carum caucasicum 1.10.1...... 0.3..0.2. Sedum tenellum 0.90.2...... 0.3..0.1. Taraxacum crepidiforme 0.80.2...... 0.3..0.1. Senecio taraxacifolius (tC Ass. 2) 1.621.51.3...... 6.1..3.1. Myosotis asiatica .1.5...... 0.20.4.0.10.20.1 Oxyria digyna .1.3...... 0.20.3.0.10.20.1 Astragalus beckii (C Ass. 3) ..1.6...... 0.4..0.2. Nepeta sahandica (C Ass. 3) ..0.9...... 0.2..0.1. Potentilla porphyrantha (C Ass. 4) ..1.12.9...... 1.0..0.5. Alopecurus dasyanthus (C Ass. 4) ...1.1...... 0.3..0.1. Potentilla argaea (C Ass. 4) ...0.6...... 0.2..0.1.

C & D Erigerontion venusti (Ev) Erigeron caucasicus subsp. venustus (C) . 2.4 1.6 1.1 ...... 1.3..0.6. Tripleurospermum caucasicum (C) 0.620.11.9...... 1.2..0.6. Festuca alaica 8.9 3.8 2.7 . . . 0.2 . . 0.2 3.9 0.1 0.1 2.0 0.1 Koeleria eriostachya 3.30.71.6...... 1.4..0.7. Galium hyrcanicum .2.21.5...... 0.9..0.5. Draba bruniifolia subsp. bruniifolia ..2.22.1...... 1.1..0.5. Sesleria phleoides .0.71.7...... 0.6..0.3. Minuartia glandulosa 1.50.80.4...... 0.7..0.3. Saxifraga sibirica 0.51.20.21.1...... 0.8..0.4. Pedicularis caucasica .0.40.9...... 0.3..0.2. Erysimum gelidum .0.50.5...... 0.3..0.1.

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C & D Ass. 5-8 Euphorbia aucheri (tC Ass. 5) ....6.32.40.1.0.2. .2.20.11.10.1 Scutellaria glechomoides (tC Ass. 5) ....5.3.0.8.0.7. .1.50.40.80.4 Nepeta racemosa (C Ass. 6) .....14.8.... .3.7.1.9. Cicer tragacanthoides (C Ass. 6) .....14.6.... .3.7.1.8. Leonurus cardiaca subsp. persicus (C Ass. 6).....1.2.... .0.3.0.2. Asperula glomerata subsp. bracteata .....0.53.1.4. .0.90.70.40.7 Senecio iranicus (C Ass. 8) ...... 3 . . . 0.1 . 0.0 . Artemisia melanolepis ...... 13 . . . 0.3 . 0.2 . Catabrosella parviflora .0.3.0.3. . .6.3 . . 0.2 0.2 . 0.2 . Carex oreophila ...... 5.3 . . . 0.1 . 0.1 . Draba siliquosa ...0.3...2.7 . 0.2 0.1 0.1 0.1 0.1 0.1 Veronica biloba 0.5...... 2.3 . . 0.1 0.1 . 0.1 . Potentilla argyroloma ...... 2 . 0.6 . 0.1 0.3 0.0 0.3

D Elymo-Astragalion macrosemii (E-Am) Astragalus macrosemius ....11.3648 . . . 3.3 . 1.6 . Elymus longearistatus ....0.54.11.8... .1.6.0.8.

C & D Physoptychio-Brometalia (P-B) Bromus tomentosus (C), (D Ass. 3 & 7) 1.1 1.7 7.2 . . 3.4 15.4 8.7 0.5 . 2.5 4.9 0.3 3.7 0.3 Ziziphora clinopodioides (C), (D Ass. 3 & 7) . . 5.9 . . 0.1 2.2 . . . 1.5 0.6 . 1.0 . Physoptychis gnaphalodes (C) . . 0.8 . 0.5 2.8 0.3 . 0.3 . 0.2 0.9 0.2 0.6 0.2 Poa araratica (D Ass. 3) . 0.1 5 0.3 . 1.1 2.5 2.7 . 0.2 1.4 1.0 0.1 1.2 0.1

C & D Ass. 9-10 Galium aucheri (C Ass. 9) ...... 2.20.2..1.2.1.2 Veronica paederotae (C Ass. 9) ...... 1.2. ..0.6.0.6 Senecio vulcanicus (C Ass. 9) ...... 0.8. ..0.4.0.4 Lamium tomentosum ...... 1.30.2..0.8.0.8 Pseudocamelina glaucophylla ...... 0.7. ..0.4.0.4 Achillea aucheri (tC Ass. 10) ...... 1 0.5 4.4 . 0.0 2.5 0.0 2.5 Veronica aucheri (C Ass. 10) ...... 0.7 . 1.2 . 0.0 0.6 0.0 0.6

C & D Didymophysion/-etalia (Da) Didymophysa aucheri (tC) . . 1.7 1.4 ....2.430.8.2.70.42.7 Dracocephalum aucheri (tC), (D Ass. 3 & 10) . 0.1 2.7 . . . 3.6 9.7 0.3 8.3 0.7 0.3 4.3 0.5 4.3 Cerastium purpurascens (C) ...... 6.3 0.8 1.3 . 0.2 1.1 0.1 1.1 Erysimum elbrusense ...... 3 0.7 0.5 . 0.1 0.6 0.0 0.6

D Didymophyso-Dracocephaletea Alopecurus textilis (D Ev) . 1.3 2.7 .....0.50.41.0.0.50.50.5

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Other species Arabis caucasica ..0.40.3....0.41.90.2.1.20.11.2 Helichrysum psychrophilum 1.4 1.6 0.4 . . 0.4 0.1 . . . 0.9 0.1 . 0.5 . Campanula stevenii subsp. beauverdiana 0.3 0.3 0.3 . . 0.1 0.2 . . . 0.2 0.1 . 0.2 . Minuartia recurva subsp. oreina ..0.2...... 0.1..0.0. Jurinella moschus ..1.6...... 0.4..0.2. Alyssum condensatum ..0.3...... 0.1..0.0. Polygonum serpyllaceum .....10.2... .0.3.0.2. Ranunculus crymophilus ...... 0.7..0.2.0.20.10.10.1 Erigeron uniflorus subsp. elbursensis ...... 0.5..0.5.0.10.30.10.3 Oxytropis persica ...... 0.2.0.2. .0.10.10.00.1 Draba pulchella .....0.20.4... .0.2.0.1. Senecio glaucus .....0.30.1... .0.1.0.1. Pisum formosum ...... 0.5. ..0.3.0.3 Alopecurus himalaicus ...... 0.2..0.1.0.1 Lepechiniella persica ...... 0.20.2..0.2.0.2 Veronica kurdica ...... 0.7 . 0.2 . 0.2 0.1 0.1 0.1 Piptatherum laterale .....0.6.... .0.2.0.1. Acantholimon demawendicum .....0.10.6... .0.2.0.1. Silene marschallii subsp. sahendica .0.5...... 0.1..0.1. Thymus kotschyanus .0.40.1...... 0.1..0.1. Scrophularia variegata subsp. variegata .0.4...0.3....0.10.1.0.1. Crepis elbursensis ..0.2...... 0.1..0.0. Vicia ciceroidea .....0.3.... .0.1.0.0. Arenaria insignis ...... 0.2... .0.1.0.0. Tanacetum polycephalum .0.1....0.3...0.00.1.0.1. Veronica orientalis 0.40.2...... 0.2..0.1. Asperula prostrata 0.3...... 0.1..0.0. Solenanthus circinnatus .....0.1.... .0.0.0.0. Astragalus sp...... 0.1.... .0.0.0.0. Thymus pubescens .....0.1.... .0.0.0.0. Semenovia frigida ...... 0.1... .0.0.0.0. Linaria lineolata .....0.1.... .0.0.0.0. Cousinia crispa .....0.1.... .0.0.0.0. Minuartia lineata .....0.1.... .0.0.0.0. Taraxacum sp...... Crepis sahendi .0.1...... 0.0..0.0. Euphorbia sahendi ..0.1...... 0.0..0.0. Tragopogon kotschyi 0.3...... 0.1..0.0. Poa bulbosa 0.3...... 0.1..0.0.

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Sibbaldia parviflora .0.1...... 0.0..0.0. Artemisia sp...... Trachydium depressum ...... 0.2. ..0.1.0.1 Draba aucheri ...... 0.3. ..0.2.0.2 Potentilla sp. . . 0.1 ...... 0.0..0.0. Kobresia schoenoides ...0.3...... 0.1..0.0. Chenopodium foliosum subsp. montanum .....0.2.2 . . . 0.6 . 0.3 .

123 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Appendix S4. Phytosociological table of the alliance Erigerontion venusti. Differential species (D) are shaded, character species (C) are additionally framed. Association numbers refer to Table 4.

Class Didymophyso aucheri-Dracocephaletea aucheri Order Physoptychio gnaphalodis-Brometalia tomentosi Alliance Erigerontion venusti Association 12 3 4 Subassociation aaaaabbbbbbcccccccccccccddddddddddd Typus of ass. (*) and subass. (°) * * ° ° * ° * 1066 1063 1064 1065 1062 1061 1060 1059 1031 1029 1027 1028 1030 1084 1083 1082 1109 1081 1073 1072 1108 1076 1075 1074 1085 1086 1087 1033 1040 1041 1039 1042 1034 1036 1035 1032 1037 1038 1106 1105 1099 1107 1096 1100 1097 1103 1102 1101 1088 1095 1093 1104 1090 1091 1089 1092 1098 1094 940 941 942 943 949 950 946 951 945 947 948 Relevé number Sabalan Sabalan Sabalan Sabalan Sabalan Sabalan Sabalan Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand Sahand

Geographical position of the plots 3353 3359 3360 3360 3358 3354 3352 3350 3451 3433 3414 3424 3476 3395 3360 3331 3663 3302 3312 3305 3680 3379 3350 3330 3450 3456 3503 3533 3519 3519 3518 3519 3520 3515 3515 3534 3513 3515 3658 3704 3703 3660 3707 3658 3624 3660 3636 3670 3658 3667 3664 3538 3622 3592 3700 3553 3567 3550 3571 3629 3709 3618 4326 4232 4350 4212 4335 4371 4357

Altitude (m a.s.l.) WNW WNW WNW WNW WNW WNW WNW WNW NNE NNE NNE ENE ENE ENE ENE ENE ENE NW NW NW NW NW NW NW NW NW NW NE NE NE NE NE NE NE NE NE NE NE W W W W W W W W W W N N N N N N N N N N N N N N N N E E E E Aspect E 35 40 40 40 35 35 35 35 40 35 30 35 40 40 35 35 45 30 35 35 35 45 40 40 45 45 45 20 30 30 35 30 25 30 30 25 30 30 15 35 30 25 25 15 30 20 30 20 15 20 20 40 30 35 30 40 45 45 45 25 20 30 15 20 15 10 15 15 15 Slope (degrees) 75 50 40 80 60 70 50 60 50 30 40 30 40 35 35 35 40 25 30 25 35 30 40 30 30 30 30 20 20 20 20 20 10 10 10 10 20 25 30 35 40 40 25 30 40 35 35 20 35 40 20 30 15 10 15 10 15 10 15 15 20 15 Cover total (%) 5 4 7 4 5 2 2 12 12 11 12 16 17 13 12 16 10 13 10 11 13 11 12 12 12 13 13 13 11 10 10 11 12 10 10 12 15 14 12 12 12 11 16 13 11 11 13 13 Species richness 8 5 5 4 8 8 8 3 3 2 7 7 6 8 7 8 8 5 8 9 9 7 9 6 9 4 2 6 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 25 25 25 25 25 25 25 25 25 25 25 Relevé area (m²) TWINSPAN classification 000000000000000000000000000000000000000000000000000000000000000000000 000000000000000000000000000111111111111111111111111111111111111111110 000000001111111111111111111000000000000000000000000000000000001111111 11111111111111111111111100000000000 111111111110000000000000 11111000000 C & D Associations (and subassoc.) Nepeta menthoides 43343433 ...... 22.211.111...... 1...... Alopecurus aucheri 2112211111211 ...... ++...... Carum caucasicum ....++1+r...... Taraxacum crepidiforme .....+++.+.r...... Sedum tenellum ....+1.+1...... Senecio taraxacifolius +1+++ .+ . .232232232222232333 ...... 11..1+11221...... Myosotis asiatica ...... 11111+ ....++.11+...... Oxyria digyna ...... +.+...1.111111...... Jurinella moschus ...... 22212 ...... Minuartia recurva subsp. oreina ...... +++...... Astragalus beckii ...... 2.21112+11...... Alyssum condensatum ...... +..1.+..+.+...... r...... Nepeta sahandica ...... +...... 111.1211...... Arabis caucasica ...... +1+.+...1.1.+..... Potentilla porphyrantha ...... 11122 ...... +...... +111111 Alopecurus dasyanthus ...... +++...+ Potentilla argaea ...... +..+...

C & D Erigerontion venusti Erigeron caucasicus subsp. venustus ...... ++1+1+11+11111111 . .+1111+++ .1+ ...... 111+++++1 ...... 11+++.+..+ Tripleurospermum caucasicum ....+.1.+1111+++1 . 11 . 111 ...... +.+1+++.+ Festuca alaica 12 .12222++12 .12111112111 ....1+11+11111..1+1+.111+11111+1+.1121...... Koeleria eriostachya + .+ .211111 . .1+1 ...... 1211++ . .++ . . .++ .+11 . .+2 ...... Galium hyrcanicum ...... ++..1121 .111 .++1 . . .+111+ .+ ...... 1+1..+..2..++.11.1...... Draba bruniifolia subsp. bruniifolia [D Subass. 3c] ...... +11.+1+1.2r2122...... +1++11+. Sesleria phleoides ...... 11.11...+...... 212..+..12..1...1...... 1...... Minuartia glandulosa [D Subass. 3b] . 1 . . +++1+ . . ++ . 1+ . . . 1 . + ...... 1.+++1+ ...... Saxifraga sibirica .....++...+.+.....1..+111++ ...... +.....+1...+++...+

Pedicularis caucasica ...... 1+..+...... +1.1...... ++.1+11+++++ ...... Erysimum gelidum ...... +11+...... +1.+1...... 1.2......

C & D Physoptychio-Brometalia tomentosi Bromus tomentosus [D Ass. 3] +.+.1+...... ++21++.11..+...21111111+122212122222222122111111+1 ...... Ziziphora clinopodioides [D Ass. 3] ...... 11+1.++1+122222111222222111+11111+1 ...... Physoptychis gnaphalodes ...... 1..+...... 11..++1+1...... ++...... Poa araratica [D Ass. 3] ...... +...... 1+++.+++1++21122212122221++ . 111 . 1++ . . . + . . .

C & D Didymophyso-Dracocephaletea aucheri Dracocephalum aucheri [D Ass. 3] ...... +...... 1112111112+ . .1+11+11 .111 .2+ ...... Didymophysa aucheri ...... 1+.11111.....11121111 . +1 . ++++ . + Alopecurus textilis [D Subass. 3c] ...... ++.++.+2..+...1...1.+....+22+3+1+11+...... 11......

Companion species Scrophularia variegata ...... 11...... +...... Helichrysum psychrophilum 1...+1.111++1+11+.+.1.1...... +...... +....+...... ++1...... Campanula stevenii subsp. beauverdiana ...... ++...+...... +...... +1++...... +...... Draba siliquosa ...... +..... Thymus kotschyanus ...... +..++...... r...... +...... Silene marschallii subsp. sahendica ...... 111...... Veronica biloba ...... ++...... Tanacetum polycephalum ...... +...... Asperula prostrata ....+...... Colpodium parviflorum ...... ++r...... +... Veronica orientalis 1...... +..+...... Potentilla argaea ...... 1...... Carex oreophila ...... +... Tragopogon kotschyi ...... +...... Poa bulbosa .....+...... Crepis sahendi ...... +...... Sibbaldia parviflora ...... +...... Crepis elbursensis ...... +...... +...+...... Taraxacum sp...... r...... Artemisia sp...... r......

124 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Appendix S5. Phytosociological table of the alliance Elymo longearistati-Astragalion macrosemii. Differential species (D) are shaded, character species (C) are additionally framed. Association numbers refer to Table 4.

Class Didymophyso aucheri-Dracocephaletea aucheri Order Physoptychio gnaphalodis-Brometalia tomentosi Alliance Elymo longearistati-Astragalion macrosemii Association 56 7 8 Subassociation a a a a aaaaaaaaaabbbbbbbaaaaaaaaaaaaaaaaaaaaaaaabbbbccccc Typus of ass. (*) and subass. (°) * * ° * 869 868 760 763 809 761 832 871 830 831 765 766 870 767 768 769 833 834 882 881 880 718 719 716 717 714 715 935 612 883 889 888 607 620 619 614 605 606 609 610 611 613 616 617 618 615 819 818 820 817 822 885 886 887 884 821 890 851 852 608 926 932 933 Relevé number Damavand Damavand Damavand Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal Tuchal

Geographical position of the plots 3235 3240 3710 3718 3718 3712 3620 3195 3619 3620 3726 3726 3200 3727 3728 3728 3625 3630 3630 3635 3640 3500 3500 3500 3500 3500 3500 3871 3900 3840 3922 3925 3967 3912 3960 3893 3965 3965 3930 3920 3904 3885 3955 3957 3950 3957 3845 3844 3868 3828 3868 3848 3850 3849 3842 3868 3930 3978 3980 3930 4223 4215 4216 Altitude (m a.s.l.) WSW SSW SSW NNE NNE ENE ENE ENE ENE ENE ENE ENE ENE ENE ESE SW SW SW SW SW SW SW SW SW SW SW SW SW SW SW SW SW NE NE N N N E E S S S S S S S S S S S E S S S S S S S S Aspect S S S S 15 15 15 25 30 15 40 15 40 40 25 25 15 25 30 30 40 35 40 45 45 40 40 40 40 40 40 35 20 20 30 30 20 20 30 30 35 35 30 30 30 30 38 35 35 33 38 10 10 35 35 27 25 25 10 15 15

Slope (degrees) 5 5 7 5 7 5 10 15 10 10 75 65 50 50 50 30 50 40 45 35 50 40 30 40 50 20 35 35 30 35 25 35 30 60 40 45 45 30 25 35 40 40 35 35 30 35 25 30 30 35 25 25 30 15 20 60 40 40 60 25 20 20 20 30 55 75 75

Cover total (%) 7 5 10 13 11 13 12 16

Species richness 2 2 2 3 3 2 7 6 7 6 8 7 4 7 7 3 7 6 9 9 7 5 6 8 5 7 8 6 6 8 9 7 8 5 5 5 4 4 4 5 5 4 5 6 3 6 5 7 4 5 8 7 4 4 2 2 3 100 100 100 100 100 100 100 Relevé area (m²) 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 25 TWINSPAN classification 111111111111111111111111111111111111111111111111111111111111111 000000000000000000000000000000000000000000000000000000000000000 000000000000000000000000000111111111111111111111111111111111111 000000111111111111111111111000000000000000000000000000000000111 111111111100100000000000000000000000011111111000011111 111111111111111100000000000011111 C & D Associations (and subassoc.) Euphorbia aucheri 1221111 .++....1...222+...... 1...... Scutellaria glechomoides . . 2211 ...... 112.1....11...... Nepeta racemosa ...... 3222 ...... 22123323232 ...... Cicer tragacanthoides ...... 333232323222 ...... Leonurus cardiaca subsp. persicus ...... +111111 . . . . . 1+...... Polygonum serpyllaceum ...... +111111 ...... 1..+...... Asperula glomerata subsp. bracteata ...... 11...+...... +.1+11+11111111122122++...... Ranunculus crymophilus ...... 2111 ...... Festuca alaica ...... 1.2...... Senecio iranicus ...... 111 Artemisia melanolepis ...... 222 Catabrosella parviflora ...... 211 Cerastium purpurascens var. elbursensis ...... 211 Erysimum elbrusense ...... 111 Draba siliquosa ...... 1+1 Veronica biloba ...... +1+ Carex oreophila ...... 2.1 Potentilla argyroloma ...... 1.1

D Elymo longearistati-Astragalion macrosemii Astragalus macrosemius 11.....1..1+1.....12...... 21322+1111112221211 .+. .+21 . 2 . . . . 1334 Elymus longearistatus . . . 1 . . 111111+111211 . 122+11+2 . 111111111+11+ ...... 1....11......

C & D Physoptychio-Brometalia tomentosi Bromus tomentosus [D Ass. 7] ...... 1.1112 . . 1 . 12121 . . 1 . 1+12222222233322222222222132222222222 . Ziziphora clinopodioides [D Ass. 7] ...... 1...... 2.11111 ...... +.121211 . . 11 ...... Physoptychis gnaphalodes ....1.....12.222..1...... +...... 1+.+...... Poa araratica ...... 1..11...... ++1+1+.+21221 ...... 1..1...... 12.2.....11+

C Didymophyso-Dracocephaletea aucheri Dracocephalum aucheri [D Subass. 7c] ...... +211..1...... 11.....1121122222221

Companion species Scrophularia variegata ...... 11...... Helichrysum psychrophilum ...... 11.+...... ++...... Campanula stevenii subsp. beauverdiana ...... 1...... 1...1+...... Arenaria insignis ...... 1.+..1...... Tanacetum polycephalum ...... 11..+.1...... Piptatherum laterale ...... +1.11..+...... Acantholimon demawendicum ...... 1...... 1+.11...+...... 11...... Achillea aucheri ...... 1 Veronica aucheri ...... + Veronica kurdica ...... + Senecio glaucus ...... +...... rr+..rr...... r...... Semenovia frigida ...... +...... Vicia ciceroidea ...... 1.....1...... Minuartia lineata ...... 1...... Solenanthus circinnatus ...... +...... Astragalus sp...... +...... Thymus pubescens ...... +...... Linaria lineolata ...... 1...... Cousinia crispa ...... +...... Erigeron uniflorus subsp. elbursensis ...... 2..1...... Oxytropis persicus ...... 11...... Draba pulchella ...... ++...... 1...... +.1+.1...... Chenopodium foliosum subsp. montanum ...... +.1...... 11

125 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Appendix S6. Phytosociological table of the alliance Didymophysion aucheri. Differential species (D) are shaded, character species (C) are additionally framed. Association numbers refer to Table 4. Relevés from Klein & Lacoste (2001) are marked in green.

Class Didymophyso aucheri-Dracocephaletea aucheri Order Didymophysetalia aucheri Alliance Didymophysion aucheri Association 9 10 Subassociation (prov.) aaaaaabbbbb Typus of associations ** 1167 1166 1171 1169 1165 1164 1168 1172 1170 1174 1173 1175 1176 1177 928 927 931 930 929 917 916 915 918 Relevé number Damavand Damavand Damavand Damavand Damavand Damavand Azadkuh Azadkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh Alamkuh

Geographical position of the plots 3850 3790 3930 3890 3780 3520 3890 3960 3900 4090 4040 4160 4250 4459 4799 4360 4395 4450 4455 4455 4455 4455 4350

Altitude (m a.s.l.) SSW SSW SSW SSE SSE SW SE SE SE W W W W N N S S S S S S S Aspect 0 30 30 25 30 33 25 30 30 30 27 20 20 25 30 17 20 25 24 Slope (degrees) 0 5 5 5 5 10 10 10 10 10 15 30 25 15 20 Cover total (%) 5 5 5 5 5 5 5 5 5 2 2 5 5 12 10 12 Species richness 9 7 7 5 5 7 5 5 6 6 9 5 4 7 5 4 4 4 5 4 200 400 100 400 200 100 100 100 100 100 100 10 10 10 10 10 10 25 25 25 25 20 Relevé area (m²) 5 TWINSPAN classification 11111111111111111111111 11111111111111111111111 00000000000011111111111 00000011111 C & D Associations (and subassoc.) Galium aucheri 1+++++ . +11+1 ...... + Veronica paederotae + . ++ . ++++ ...... Senecio vulcanicus ++.+...++...... Lamium tomentosum ....+++++1+ . + ...... Asperula glomerata subsp. bracteata .1+.+1+1.+...... Pseudocamelina glaucophylla +++..+...... Achillea aucheri +..+...... +.111222 ..... Veronica aucheri ...... +1+++....+ Potentilla argyroloma ...... +.1.+ Arabis caucasica ...... 1+...... 112+

C Didymophysion aucheri Cerastium purpurascens var. elbursensis ..+..+..+..1...+..111.1

C & D Didymophysetalia aucheri Didymophysa aucheri +. 11+1111+1+11111111111 Dracocephalum aucheri ...... 1....++.+113222+ Erysimum elbrusense ...... ++.+++r.1......

C & D Physoptychio-Brometalia tomentosi Bromus tomentosus ++....+...... Physoptychis gnaphalodes ....1...... Poa araratica ...... +...... Festuca alaica ...... +...... Scutellaria glechomoides ...... ++++ ...... Euphorbia aucheri ...... +......

D Didymophyso-Dracocephaletea aucheri Alopecurus textilis .+.....+....+...... +

Companion species Pisum formosum 11...... Trachydium depressum ..+...... Draba aucheri ...... 1...... Lepechiniella persica ...... +...+...... Oxytropis persicus ...... +...... Alopecurus cf. himalaicus ...... +...... Draba siliquosa ...... +...... Erigeron uniflorus subsp. elbursensis ...... 1.....+ Veronica kurdica ...... +. Myosotis asiatica ...... + Oxyria digyna ...... + Ranunculus crymophilus ...... +

126 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3 A

Photo S1. Character species of the class: A) Didymophysa aucheri in association of Dracocephalo-Brometum tomentosi (Sahand Mt. 3570 m a.s.l., photo: J. Noroozi), B) Dracocephalum aucheri (Alamkuh Mt. 4200 m a.s.l., photo: J. Noroozi). 127 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Photo S2. Nepetetum menthoidis (Sahand Mt. 3360 m a.s.l., photo: J. Noroozi): Nepeta menthoides.

Photo S3. Senecionetum taraxacifolii (Sahand Mt. 3550 m a.s.l., photo: J. Noroozi): Senecio taraxacifolius.

128 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Photo S4. Dracocephalo aucheri-Brometum tomentosi jurinelletosum moschi (Sahand Mt. 3520 m a.s.l., photo: J. Noroozi): Jurinella moschus.

Photo S5. Dracocephalo aucheri-Brometum tomentosi minuartietosum glandulosae (Sahand Mt. 3515 m a.s.l., photo: J. Noroozi): Dracocephalum aucheri.

129 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Photo S6. Dracocephalo aucheri-Brometum tomentosi typicum (Sahand Mt. 3630 m a.s.l., photo: J. Noroozi): Dracocephalum aucheri, Bromus tomentosus, Festuca alaica.

Photo S7. Dracocephalo aucheri-Brometum tomentosi nepetosum sahandicae (Sahand Mt. 3590 m a.s.l., photo: J. Noroozi), Nepeta sahandica.

130 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Chapter 3

Photo S8. Potentilletum porphyranthae (Sabalan Mt. 4300 m a.s.l., photo: J. Noroozi): Erigeron caucasicus subsp. venustus, Potentilla porphyrantha, Draba bruniifolia subsp. bruniifolia.

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2 1

B C

Photo S9. A) Scutellario glechomoidis-Euphorbietum aucheri [1] is surrounded with Cicero tragacanthoidis-Nepetetum racemosae [2] (Tuchal Mt. 3710 m a.s.l., photo: J. Noroozi), B) Scutellaria glechomoides (photo: J. Noroozi), C) Euphorbia aucheri with long and flexible rhizomes between the screes (photo: J. Noroozi).

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Photo S10. Cicero tragacanthoidis-Nepetetum racemosae typicum (Tuchal Mt. 3600 m a.s.l., photo: J. Noroozi): Cicer tragacanthoides, Nepeta racemosa, Leonurus cardiaca subsp. persicus.

Photo S11. Cicero tragacanthoidis-Nepetetum racemosae polygonetosum serpyllacei (Tuchal Mt. 3640 m a.s.l., photo: J. Noroozi): Nepeta racemosa.

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Photo S12. Asperulo bracteatae-Brometum tomentosi typicum (Tuchal Mt. 3900 m a.s.l., photo: J. Noroozi): Bromus tomentosus, Astragalus macrosemius.

Photo S13. Asperulo bracteatae-Brometum tomentosi (Tuchal Mt. 3880 m a.s.l., photo: J. Noroozi): Dracocephalum aucheri, Bromus tomentosus.

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Photo S14. Senecio iranici-Astragaletum macrosemii (Damavand Mt. 4210 m a.s.l., photo: J. Noroozi): Astragalus macrocemius, Dracocephalum aucheri, Senecio iranicus.

Photo S15. Galietum aucheri Klein & Lacoste 1999 (Alamkuh Mt. 4200 m a.s.l., photo: J. Noroozi): Veronica paederotae.

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Photo S16. Dracocephaletum aucheri achileetosum aucheri (Damavand Mt. 4470 m a.s.l., photo: J. Noroozi): Achillea aucheri.

Photo S17. Dracocephaletum aucheri typicum (Alamkuh Mt. 4455 m a.s.l., photo: J. Noroozi): Cerastium purpurascens var. elbursensis, Dracocephalum aucheri.

136 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Conclusions

Conclusions

With this thesis, our knowledge about the phytodiversity, phytogeography, ecology and phytosociology of the high elevations of Iran markedly increased, and data of high-elevation vegetation in Iran has multiplied and are available as basic references for the future studies. For the first time, the papers presented in this thesis provide a country-wide overview of the vertical and geographical distribution patterns of vascular plant species living in the uppermost elevation zones in Iran and the the syntaxonomic position of the scree vegetation of the high elevations of Iran has been clarified and compared with the vegetation of similar habitats in neighboring mountain regions and in the Alps.

The main conclusions are summarized as follows: 1) The vascular plant flora of subnival-nival habitats of Iran is not outstandingly rich in species, however, the rate of endemism and narrowly distributed species are very high. The outstanding level of high-altitude endemism appears to result mainly from a pronounced orographic isolation of the country’s highly scattered and fragmented cold areas and from the absence of extensive Pleistocene glaciations. Glaciers were present mainly in the uppers valleys, whereas high-elevation mountain slopes commonly remained ice-free therefore could have functioned as long-term refugia of cold-adapted plant species. 2) Several vascular plant species, restricted to the subnival-nival belt, were thus far unknown to science and other species were previously only known from mountains outside of Iran. This led to new original descriptions of species and the revision of disjunct distribution patterns of some species. We, hence, expect that the upper elevation zones of Iranian mountains are underexplored and that further unknown species occur. Furthermore, our knowledge about the habitats of rare species in different mountains is low. Therefore, we highly recommend more field exploration in high mountains to determine the ecology, biology and threatened rank of local endemics according to IUCN criteria, and improve the protection status of the high mountain flora and vegetation. 3) Our vegetation study extends knowledge of alpine and subnival scree vegetation into a previously almost unknown area, and therefore represents an important building block for a future synopsis of the scree vegetation of the northern hemisphere. It is a basis for monitoring of future

137 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Conclusions

changes and for assessing endangered vegetation types and for assessing nature conservation needs. 4) One of the most important threats to the flora and vegetation of the study habitats can be the predicted climate change, which may seriously shrink the size of suitable habitats of the unique subnival–nival flora of Iran. The potential of alternative low-temperature habitats at higher elevations is very limited and, therefore, the survival of high-elevation species is uncertain. Decreasing water availability during the growing season and land use pressure (Kousari et al. 2011; Zarghami et al. 2011) may further exacerbate the situation. Additional pressure through drought-tolerant competitors from lower altitudes can increase the risk of biodiversity losses. A decline of high-altitude habitats through climate warming and reduced water availability is expected to be similar to that on Mediterranean mountains of S Europe, where species richness has decreased during the past particularly warm decade (Pauli et al. 2012). Hence, we have to expect a very high risk of biodiversity losses in the high-altitude vegetation of N and NW Iran. Detailed knowledge of these plant assemblages appears to be essential for tracing the expected changes in their species composition and the possible shrinkage of cryophilic, narrowly distributed species. 5) One of the major aims of this PhD thesis was assessing the effects of climate changes on alpine habitats of Iran. Two monitoring stations, therefore, were established in Central Alborz and in Sahand (NW Iran) according to the GLORIA approach. The duration of my PhD studies, however, was too short to allow for a resurvey of the vegetation recorded in the permanent plots. The collected permanent plot data are uploaded to the central GLORIA database and further field recording in the next years is planned to complete the Sahand site. Both Iranian GLORIA sites are planned to be resurveyed at intervals of 5 to 10 years. 6) Overgrazing is another major threat to the flora and vegetation of high elevations in Iran. The alpine grasslands are frequently used for mountain pasturing with large herds of sheep and goats during the summer growing season, when grazing grounds at lower elevations fall dry. Overgrazing and trampling of the herds threat the fragile communities of alpine and even subnival zones. In spite of the low attraction of subnival zones for sheep and goats to feed, effects of pastoralism should not be neglected, especially the impact of trampling on unstable scree habitats. Large herds cause substrate movement, plants can be uprooted and thus the vegetation may be completely destroyed at some sites.

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7) As my thesis mainly focuses on the alpine to subnival-nival vegetation of Alborz and NW Iran, large high-altitude areas still remain unstudied. For a comprehensive description of the country’s alpine and subnival-nival vegetation types, studies in vegetation ecology and phytosociology in the extensive Zagros mountain range appear to be indispensable.

References

Pauli, H., Gottfried, M., Dullinger, S., Abdaladze, O., Akhalkatsi, M., Benito Alonso, J.L., Coldea, G., Dick, J., Erschbamer, B., Fernández Calzado, R., Ghosn, D., Holten, J.I., Kanka, R., Kazakis, G., Kollár, J., Larsson, P., Moiseev, P., Moiseev, D., Molau, U., Molero Mesa, J., Nagy, L., Pelino, G., Puşcaş, M., Rossi, G., Stanisci, A., Syverhuset, A.O., Theurillat, J.-P., Tomaselli, M., Unterluggauer, P., Villar, L., Vittoz, P. & Grabherr, G. (2012) Recent plant diversity changes on Europe’s mountain summits. Science 336: 353-355. Kousari, M.R., Ekhtesasi M.R., Tazeh M., Saremi Naeini, M.A. & Asadi Zarch, M.A. (2011) An investigation of the Iranian climatic changes by considering the precipitation, temperature, and relative humidity parameters. Theoretical Applied Climatology 103:321–335. Zarghami, M., Abdi, A., Babaeian, I., Hassanzadeh, Y. & Kanani, R. (2011) Impacts of climate change on runoffs in East Azerbaijan, Iran. Global and Planetary Change 78: 137–146.

139 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Curriculum Vitae

Curriculum Vitae Jalil Noroozi Eshlaghi

Academic education: - 1998-2002: BSc in Plant Biology, Department of Plant Science, Faculty of Natural Science, University of Tabriz, Tabriz, IRAN. - 2002-2005: MSc in Plant Biology (Ecology and Systematic), Department of Biology, Faculty of Science, University of Tehran, Tehran, IRAN. - 2009-2013: PhD study in Plant Ecology, Department of Conservation Biology, Vegetation and Landscape Ecology, University of Vienna, AUSTRIA.

Main Research Interests: - Ecology of alpine habitats - Biogeography and evolution of alpine plants - Alpine plant communities in Iran and the Caucasus region - Alpine plant diversity and climate change - Land use impacts and conservation biology in alpine environments

MSc Thesis Subject: Phytosociological study of alpine zone of Tuchal Mountains, N. Tehran. University of Tehran, Iran.

PhD Thesis Subject: Comparative analysis of high-alpine and subnival vegetation of mountain ecosystems in Iran (Alborz and NW-Iran) and assessing the impacts of climate change and land-use

Publications: Book: - Noroozi, J. (2010) A glance at the wild flowers of Iranian mountains. Karimkhaneh Zand Publication, Tehran, IR. Book Chapters: - Noroozi, J. (In press). High mountain regions of Iran (chapter 7.6), in C. Hobohm (editor). Endemism in Vascular plants. Springer. - Noroozi, J. (In press). Iran (chapter 2), in J. Solomon, T. Shulkina & G. Schatz (editors). Red Book of endemic plants of the Caucasus Region. Missouri Botanical Garden Press, Saint Louis.

140 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Curriculum Vitae

International Peer-reviewed Journals: - Noroozi, J., Willner, W., Pauli, H. & Grabherr, G. (2013) Phytosociology and ecology of the high-alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.). Applied Vegetation Science DOI: 10.1111/avsc.12031. - Butterfield, B.J., Cavieres, L.A., Callaway, R.M., Cook, B.J., Kikvidze, Z., Lortie, C.J., Michalet, R., Pugnaire, F.I., Schöb, C., Xiao, S., Anthelme, F., Björk, R.G., Dickinson, K., Gavilán, R., Kanka, R., Maalouf, J.-P., Noroozi, J., Parajuli, R., Phoenix, G.K., Reid, A., Ridenour, W., Rixen, C., Wipf, S., Zhao, L., Brooker, R.W. (2013) Alpine cushion plants inhibit the loss of phylogenetic diversity in severe environments. Ecology Letters DOI: 10.1111/ele.12070. - Noroozi, J., Ajani, Y. (2013) A new alpine species of Nepeta sect. Capituliferae (Labiatae) from Northwestern Iran. Novon 22: 297–303. - Akhani, H., Mahdavi, P., Noroozi, J. & Zarrinpour, V. (2013) Vegetation patterns of the Irano-Turanian steppe along a 3,000 m altitudinal gradient in the Alborz Mountains of Northern Iran. Folia Geobotanica 48: 229-255. - Noroozi, J., Pauli, H., Grabherr, G. & Breckle, S.W. (2011) The subnival-nival vascular plant species of Iran: a unique high-mountain flora and its threat from climate warming. Biodiversity and Conservation 20: 1319-1338. - Noroozi, J., Akhani, H. & Willner, W. (2010) Phytosociological and ecological study of the high alpine vegetation of Tuchal Mountains (Central Alborz, Iran). Phytocoenologia 40: 293- 321. - Ajani, Y., Noroozi, J. & Levichev, I.G. (2010) Gagea alexii (Liliaceae), a new record from subnival zone of southern Iran with key and notes on sect. Incrustatae. Pakistan Journal of Botany 42: 67-77. - Noroozi, J., Ajani, Y., Nordenstam, B. (2010) A new annual species of Senecio (Compositae- Senecioneae) from subnival zone of southern Iran with comments on phytogeographical aspects of the area. Compositae News Letter 48: 43-62. - Noroozi, J. (2009) Understanding the effects of climate change on alpine biodiversity of Alborz mountains. BC Grasslands, Temperate Grasslands of the World, winter 2008/2009. 15- 17. - Noroozi, J., Akhani, H. & Breckle, S.W. (2008) Biodiversity and phytogeography of the alpine flora of Iran. Biodiversity and Conservation 17: 493-521. - Khassanov F., Noroozi J. & Akhani H. (2006) Two new species of Allium genus from Iran. Rostaniha 7(2): 119-130.

Participation at scientific meetings: - Noroozi, J. Alpine flora and vegetation of Iranian mountains and their sensitivity to global warming. Plant Life of SW Asia 8 conference at the Royal Botanic Garden Edinburgh. Edinburgh, Scotland/UK, 1-5th July 2013. (oral presentation) - Noroozi, J., Willner, W., Pauli, H., Grabherr, G. Phytosociology and ecology of the high alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.), 56th

141 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Curriculum Vitae

Symposium of the International Association for Vegetation Science (IAVS). Tartu, Estonia, 26- 30th June 2013. (poster presentation) - Noroozi, J. Alpine flora and vegetation of Iranian mountains and their sensitivity to global warming. Plant Life of SW Asia 8 conference at the Royal Botanic Garden Edinburgh. Edinburgh, UK, 1-5th July 2013. (oral presentation) - Noroozi, J., Willner, W., Pauli, H., Grabherr, G. Phytosociology and ecology of the high alpine to subnival scree vegetation of N and NW Iran (Alborz and Azerbaijan Mts.). 7th Annual Meeting of the Specialist Group on Macroecology of the Gesellschaft für Ökologie. Göttingen, Germany, 13-15th March 2013. (oral presentation) - Noroozi, J. The alpine flora of Iranian mountains and its sensitivity to climate change. International Conference on Managing Protected Areas under Climate Change – IMPACT. Dresden, Germany, 24-26 th September 2012. (oral presentation) - Noroozi, J. An overview of alpine vegetation of Iranian mountains. 21st Workshop of European Vegetation Survey. Vienna, Austria, 24-27th May 2012. (oral presentation) - Noroozi, J., Pauli, H., Grabherr, G. & Breckle S.W. Refugia for coldadapted species in the highly fragmented subnival –nival habitats of Iran. Annual Conference of the Society for Tropical Ecology (gtoe) “Islands in lands- and sea-scape: the challenge of fragmentation”. Erlangen, Germany, 22-25th February 2012. (oral presentation) - Noroozi, J., Willner, W., Pauli, H. & Grabherr, G. Phytosociological and ecological studies of high alpine to subnival scree vegetations of N and NW of Iran (Alborz and Azarbayjan Mts.). 20st Workshop of European Vegetation Survey. Rome, Italy, 06-09th April 2011. (oral presentation) - Noroozi, J., Pauli, H. & Grabherr, G. Potential effects of climate warming on a unique cryo- flora: the endemic high alpine to subnival plants in the mountains of Iran. International conference of global change and the world's mountains. Perth, Scotland/UK, 26-30th September 2010. (oral presentation) - Noroozi, J., Pauli, H. & Grabherr, G. Potential effects of climate warming on a unique high- altitude flora. 5th International GLORIA conference. Perth, Scotland/UK, 23-25th September 2010. (oral presentation) - Noroozi, J., Pauli, H. & Grabherr, G. Potential effects of climate warming on a unique cryo- flora: the endemic high alpine to subnival plants in the mountains of Iran. International GMBA-DIVERSITAS Conference on “Functional Significance of Mountain Biodiversity”. Chandolin (Valais), Switzerland, 27-30th July 2010. (oral presentation) - Noroozi, J. Caucasian Part of Iran. 3th Workshop of Coordination and Development of Plant Red List Assessments for the Caucasus Biodiversity Hotspot, CEPF-IUCN. Tbilisi, Georgia. 25-28th May 2009. (oral presentation) - Noroozi, J. Endemic species of Caucasian territory of Iran. International Symposium for Plant Conservation in the Caucasus Region. 2-8th October 2009. Missouri Botanical Garden, Missouri, USA. (oral presentation) - Noroozi, J. Alpine Grasslands of Iran. World Temperate Grasslands Conservation Initiative Workshop: Planning for a Decade of Progress. Hohhot, China, 28-29th June 2008. (oral presentation)

142 PhD Thesis of Jalil Noroozi Eshlaghi 2013 Curriculum Vitae

- Noroozi, J. XXI International Grasslands Congress/ VIII International Rangelands Congress (IGC/IRC). Hohhot, China, 29th June -5 July 2008. (oral presentation) - Noroozi, J. & Akhani, H. Biodiversity of Alpine Flora of Iran. XVII International Botanical Congress. Vienna, Austria, 17-23 th July 2005. (poster presentation)

Invited Talks: - Flora and vegetation of alpine habitats of Iran. Institut für Pflanzenwissenschaften -Botanischer Garten-, Karl-Franzens-Universität Graz, Graz, Austria. 27th March 2012. - Flora und Vegetation der alpinen Stufe im Iran. Botanical Society of Regensburg, University of Regensburg, Germany. 7th November 2011. - Flora und Vegetation der alpinen Stufe im Iran. Institute of Botany, University of Basel, Switzerland. 29th October 2010. - Mountain vegetation types of Caucasus region of Iran. Faculty of Natural Sceince, University of Tabriz, Iran. 15th December 2009. - Alpine habitats of Iranian mountains. Faculty Centre of Biodiversity, University of Vienna, Austria. 25th September 2009.

Memberships of scientific networks and societies: - Global Observation Research Initiative in Alpine Environments (GLORIA) - Temperate Grasslands Conservation Initiative (TGCI)_IUCN-WCPA - Alpine Garden Society (AGS) - International Association for Vegetation Science (IAVS) - European Vegetation Survey (EVS)

Language Skills: Farsi (Persian): native Turkish: native English: fluently (orally and written) German: good (orally), moderate (written) Arabic: basic

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